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Dietary lipid quality affects temperature-mediated reaction norms of a freshwater key herbivore
(2012)
Temperature-mediated plasticity in life history traits strongly affects the capability of ectotherms to cope with changing environmental temperatures. We hypothesised that temperature-mediated reaction norms of ectotherms are constrained by the availability of essential dietary lipids, i.e. polyunsaturated fatty acids (PUFA) and sterols, as these lipids are involved in the homeoviscous adaptation of biological membranes to changing temperatures. A life history experiment was conducted in which the freshwater herbivore Daphnia magna was raised at four different temperatures (10, 15, 20, 25A degrees C) with food sources differing in their PUFA and sterol composition. Somatic growth rates increased significantly with increasing temperature, but differences among food sources were obtained only at 10A degrees C at which animals grew better on PUFA-rich diets than on PUFA-deficient diets. PUFA-rich food sources resulted in significantly higher population growth rates at 10A degrees C than PUFA-deficient food, and the optimum temperature for offspring production was clearly shifted towards colder temperatures with an increased availability of dietary PUFA. Supplementation of PUFA-deficient food with single PUFA enabled the production of viable offspring and significantly increased population growth rates at 10A degrees C, indicating that dietary PUFA are crucial for the acclimation to cold temperatures. In contrast, cumulative numbers of viable offspring increased significantly upon cholesterol supplementation at 25A degrees C and the optimum temperature for offspring production was shifted towards warmer temperatures, implying that sterol requirements increase with temperature. In conclusion, essential dietary lipids significantly affect temperature-mediated reaction norms of ectotherms and thus temperature-mediated plasticity in life history traits is subject to strong food quality constraints.
The understanding of environmentally induced changes in the biochemical composition of phytoplankton species is of great importance in both physiological studies and ecological food web research. In extensive laboratory experiments we tested the influence of two different temperatures (10 degrees C and 25 degrees C) and a phosphorus supply gradient on the sterol concentrations of the three freshwater phytoplankton species Scenedesmus quadricauda, Cryptomonas ovata and Cyclotella meneghiniana. The diatom C. meneghiniana was additionally exposed to a silicate gradient. In two separate experiments we analysed (1) possible interactive effects of temperature and phosphorus supply and (2) the effect of four phosphorus levels and three silicate levels on algal sterol concentrations. We observed that sterol concentrations were higher at 25 degrees C than at 10 degrees C in S. quadricauda and C. meneghiniana, but were not affected by temperature in C. ovata. Interactive effects of temperature and phosphorus supply on sterol concentrations were found in C. meneghiniana. This presumably was due to the bioconversion of one sterol (24-methylenecholesterol) into another (22-dihydrobrassicasterol). Increasing phosphorus supply resulted in species-specific effects on sterol concentrations, viz. an optimum curve response in S. quadricauda, a saturation curve response in C. meneghiniana and no change in sterol concentration in C. ovata. Effects of silicate supply on the sterols of C. meneghiniana equalled the effects of phosphorus supply. Albeit we did not observe a general trend in the three phytoplankton species tested, we conclude that sterol concentrations of phytoplankton are strongly affected by temperature and nutrient supply. Interactive effects point out the importance of taking into account more than just one environmental factor when assessing the effects of environmentally induced changes on phytoplankton sterol concentrations.
There is growing consensus that the growth of herbivorous consumers is frequently limited by more than one nutrient simultaneously. This understanding, however, is based primarily on theoretical considerations and the applicability of existing concepts of co-limitation has rarely been tested experimentally. Here, we assessed the suitability of two contrasting concepts of resource limitation, i.e. Liebigs minimum rule and the multiple limitation hypothesis, to describe nutrient-dependent growth responses of a freshwater herbivore (Daphnia magna) in a system with two potentially limiting nutrients (cholesterol and eicosapentaenoic acid). The results indicated that these essential nutrients interact, and do not strictly follow Liebigs minimum rule, which consistently overestimates growth at co-limiting conditions and thus is not applicable to describe multiple nutrient limitation of herbivorous consumers. We infer that the outcome of resource-based modelling approaches assessing herbivore population dynamics strongly depends on the applied concept of co-limitation.
1. Herbivores show stronger control of element homoeostasis than primary producers, which can lead to constraints in carbon and nutrient transfer efficiencies from plants to animals. Insufficient dietary phosphorus (P) availability can cause reduced body P contents along with lower growth rates of animals, leading to a positive relationship between growth and body P.
2. We examined how a second limiting food component in combination with dietary P limitation influences growth and P homoeostasis of a herbivore and how this colimitation influences the hypothesized positive correlation between body P content and growth rates. Therefore, we investigated the responses in somatic growth and P stoichiometry of Daphnia magna raised on a range of diets with different amounts of P and the sterol cholesterol.
3. Somatic growth rates of D. magna increased asymptotically with increasing P as well as with increasing cholesterol availability. The body P content increased with increasing dietary P and stabilized at high dietary P availability. The observed plasticity in D. magna's P stoichiometry became stronger with increasing cholesterol availability, i.e. with decreasing colimitation by cholesterol.
4. At P-limiting conditions, the positive correlation between body P content and growth rate, as predicted by the growth rate hypothesis (GRH) applied to the within-species level, declined with increasing cholesterol limitation and disappeared entirely when cholesterol was not supplied. Thus, even when Daphnia shows no growth response owing to strong limitation by the colimiting nutrient, the body P content may vary substantially, calling into question the unconditional use of herbivores' P content as predictor of a potential P limitation in nature.
5. The observed interaction between dietary P and cholesterol on Daphnia's growth and stoichiometry can be used as a conceptual framework of how colimiting essential nutrients affect herbivore homoeostasis, and provide further insights into the applicability of the GRH within a consumer species.
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.
We present data on eicosapentaenoic acid (EPA)-limited growth responses of Daphnia magna under different temperatures and different dietary cholesterol availabilities to assess how EPA growth saturation thresholds depend on changing environmental conditions. D. magna was raised on gradients of dietary EPA at 15 degrees C and 20 degrees C with high cholesterol supply and at 20 degrees C with low and high cholesterol supply in laboratory experiments. A new method was applied to estimate EPA growth saturation thresholds on the basis of fitted saturation curves using bootstrapped data. The EPA threshold at which 75% and 90% of maximum growth was reached ranged from 0.7 to 1.6 mu g EPA (mg dietary C)(-1) and 2.0 to 4.9 mu g EPA (mg dietary C)(-1), respectively. Previously reported EPA concentrations in natural seston of many different lakes suggest that the thresholds measured here indicate a frequent potential for at least moderate EPA limitation in nature. Furthermore, the calculated EPA thresholds were higher in treatments of low compared with high temperature and higher in treatments of low compared with high cholesterol availability. The EPA-dependent growth responses were more strongly affected by temperature than by cholesterol availability. Our results suggest that EPA growth saturation thresholds for a particular Daphnia species probably vary in nature under different environmental conditions.
Carbon assimilation mode in mixotrophs and the fatty acid composition of their rotifer consumers
(2009)
P>1. We examined an important ecophysiological link between the mixotrophic flagellate Chlamydomonas acidophila and its consumers, the rotifers Elosa worallii, Cephalodella sp. and Brachionus sericus, by comparing their fatty acid profiles. 2. The mixotrophic flagellate was grown under either exclusively autotrophic conditions in the light, under exclusively heterotrophic conditions in the dark with an organic carbon source (glucose), or in the light plus the organic carbon sources (=mixotrophic). 3. Under heterotrophic growth conditions, C. acidophila strongly reduced its content of the n-3 polyunsaturated fatty acid (PUFA) alpha-linolenic acid (ALA, C18:3n-3) compared with auto- and mixotrophic growth conditions. Although PUFAs with more than 18 carbon atoms were not detected in C. acidophila, significant amounts of eicosatetraenoic (ETA, 20:4n-3) and eicosapentaenoic acid (EPA, 20:5n-3) were found in three rotifer consumers. 4. Species-specific differences in the fatty acid profiles with respect to ETA, EPA and the precursor ALA were found in the rotifers: Brachionus and Cephalodella fed on the heterotrophic diets synthesised less EPA. In Elosa, smaller amounts of ALA were detected but were converted efficiently to a constant content of EPA and to an exceptionally high content of ETA. 5. Since in nature the mode of carbon assimilation among mixotrophic organisms differs, and their fatty acid composition varies depending on their mode of carbon assimilation, the availability of ALA might be critical for their consumers. An insufficient dietary supply of this precursor for the synthesis of ETA and EPA can prevent consumers from regulating their content of ETA and EPA. Therefore, observed differences in values of the latter might underly species-specific differences in the competitive capability of consumers.
Enhanced water temperatures promote the occurrence of cyanobacterial blooms, which may be detrimental to aquatic herbivores. Especially, the often-dominant crustaceans could be negatively affected because cyanobacteria are deficient in phytosterols, which are required by the crustaceans to form the membrane component cholesterol, which in turn plays a role in thermal adaptation. Here, we determined the influence of temperature on growth, reproduction and the allocation of dietary sterol into somatic tissues and eggs of the keystone species Daphnia magna raised along a dietary cholesterol gradient. Mass-specific growth rates of D. magna increased with the increasing availability of dietary cholesterol up to an incipient limiting level, which increased with increasing temperature. This indicates a higher demand for cholesterol for growth at higher temperatures and may explain the consistently smaller clutch sizes of reproducing females at the highest temperature. The cholesterol content of the individuals increased with increasing dietary cholesterol; this increase was enhanced at higher temperatures, indicating a higher demand for cholesterol for tissues and probably specifically for membranes. Surprisingly, the daphnids showed different allocation strategies with regard to temperature and dietary sterol availability. The cholesterol content of eggs was enhanced at higher temperature, which suggested that females allocate more cholesterol to their offspring, presumably to ensure sufficient egg development. When dietary cholesterol was limiting, however, females did not allocate more cholesterol to their eggs. Our data suggest that during cyanobacterial blooms, a potential dietary sterol limitation of Daphnia can be intensified at higher water temperatures, which can occur with global warming.
Empirical data providing evidence for a colimitation of an herbivore by two or more essential nutrients are scarce, particularly in regard to biochemical resources. Here, a graphical model is presented, which describes the growth of an herbivore in a system with two potentially limiting resources. To verify this model, life-history experiments were conducted with the herbivore Daphnia magna feeding on the picocyanobacterium Synechococcus elongatus, which was supplemented with increasing amounts of cholesterol either in the presence or the absence of saturating amounts of eicosapentaenoic acid (EPA). For comparison, D. magna was raised on diets containing different proportions of S. elongatus and the cholesterol- and EPA-rich eukaryotic alga Nannochloropsis limnetica. Somatic and population growth of D. magna on a sterol- and EPA-deficient diet was initially constrained by the absence of sterols. With increased sterol availability, a colimitation by EPA became apparent and when the sterol requirements were met, the growth- limiting factor was shifted from a limitation by sterols to a limitation by EPA. These data imply that herbivores are frequently limited by two or more essential nutrients simultaneously. Hence, the concept of colimitation has to be incorporated into models assessing nutrient-limited growth kinetics of herbivores to accurately predict demographic changes and population dynamics.
The mode of nutrition of mixotrophic flagellates determines the food quality for their consumers
(2007)
Animals depend on a large set of essential compounds in their food. A reduced supply of specific lipid compounds has been shown to be critical for growth, reproduction and survival of aquatic molluscs and might also be critical for terrestrial gastropods. The effects were investigated of different fatty acid and cholesterol supplies on the growth, reproduction, and survival of the land snail Arianta arbustorum. Surprisingly, differences in diet did not affect shell growth, time to reach adulthood, and reproductive traits such as the number of eggs, dry mass and volume of spermatophores. However, snails reared on low cholesterol diets and low polyunsaturated fatty acid (PUFA) diets decreased their consumption rates and had a higher mortality. Snails fed a PUFA-rich diet showed higher mating activity than snails fed a diet deficient in PUFAs. A depletion of internal PUFA reserves may affect the regulation of mating behaviour because PUFAs play an important role as precursors for signal-transduction involved in the regulation of mating and reproduction. In contrast, when the diets were deficient in cholesterol, mating activity was not affected
The absence of essential biochemical nutrients, such as polyunsaturated fatty acids or sterols, has been considered as a mechanism determining trophic interactions between the herbivore Daphnia and its phytoplankton food source. Here, we experimentally quantify the sensitivity of two Daphnia species to decreasing amounts of dietary sterols by measuring variations in life history traits. The two species Daphnia magna and D. galeata were fed different mixtures of the sterol-containing green alga Scenedesmus obliquus and the sterol-free cyanobacterium Synechococcus elongatus; a higher proportion of Synechococcus in the food is equivalent to a decrease in dietary sterols. To address the significance of sterol limitation, the Daphnia species were also fed Synechococcus supplemented with cholesterol. In both species, somatic and population growth rates, maternal dry mass, the number of viable offspring, and the probability of survival were significantly reduced with the lower availability of sterols. A high correlation between the sterol content of the mixed diet and the somatic and population growth rates was found, and growth on cholesterol- supplemented Synechococcus fitted well into this correlation. Somatic growth of first-clutch neonates grown on 100% Synechococcus exhibited a pattern similar to that of somatic growth of their mothers grown on the different food regimes, which demonstrated the significance of maternal effects for sterol-limited population growth. Daphnia galeata had a twofold higher incipient limiting sterol level than D. magna, which indicated interspecific differences in sterol requirements between the two Daphnia species. The results suggest a strong impact of dietary sterols on life history traits and therefore, population dynamics of the keystone species Daphnia
Food quality controls egg quality of the zebra mussel Dreissena polymorpha : The role of fatty acids
(2004)
We investigated the investment of adult Dreissena polymorpha to the eggs by sampling mussels monthly from 4- and 15-m water depth. The fatty acid composition of eggs differed significantly between depths and over time. To assess whether temperature and food conditions led to the differences observed for mussels sampled from the two depths, mussels were reared in the laboratory under two different 3-month temperature regimes, simulating the temperature of the lake at 4- and 15-m depth. Possible effects of food quality were tested in each simulation using four diets differing in fatty acid composition: Cryptomonas erosa, Nannochloropsis limnetica [rich in polyunsaturated fatty acids (PUFAs) and long- chained PUFAs (.C18)], Scenedesmus obliquus, and the cyanobacterium Aphanothece sp. (deficient in long-chained PUFAs). In newly released eggs, specific (n-3) and (n-6) long-chained PUFAs increased when these fatty acids were available in the natural seston or in the laboratory diets. Mussels fed organisms deficient in long-chained PUFAs were still able to allocate arachidonic acid, eicosapentaenoic acid, and docosahexaenoic acid into eggs, which suggests that minimum levels of particular fatty acids were maintained in eggs by transfer from internal reserves of the female mussels to oocytes. In contrast to the diet, there were no effects of the temperature on the fatty acid composition of eggs.
Animals depend on a large set of essential compounds in their food. However, not all units of food are equal from a nutritional point of view. A reduced supply of protein and calcium might be critical for the growth, reproduction and survival of herbivorous gastropods. We experimentally examined the effects of different protein and calcium supplies on the growth and survival of the land snail Arianta arbustorum. Groups of snails were reared on agar-based diets with each of three levels of protein and calcium (nine treatments). Snails fed a high-protein diet grew faster and reached adulthood earlier and at a larger adult size than snails fed intermediate and low-protein diets. Surprisingly, the calcium concentration did not affect shell growth and adult size. Snails reared on intermediate- and low-calcium diets increased their consumption rates, but, despite this compensatory feeding, these snails were unable to take up the amount of calcium required for metabolism and shell growth and had a higher mortality. The calcium deficiency could partly be mitigated by the snails' internal calcium storage and reallocation capacity. A depletion of internal calcium reserves adversely affects the entire nutrient metabolism and was the probable cause for the high mortality rates associated with the intermediate- and low calcium diets observed in the present study
Species such as Dreissena polymorpha sometimes contribute substantially in the transfer of primary to secondary production. During the ontogenetic cycle, the reproductive investment of adult mussels is one of the main parameters that affect recruitment success. We studied how food quality and temperature affect the reproductive investment in term of egg mass of D. polymorpha in a lake by sampling mussels monthly from 4 m and 15 m depths. Temperature affected reproduction directly and also indirectly through the food. To assess whether temperature and/or food conditions led to the differences observed in mussels sampled from the two depths, mussels were reared in the laboratory under two different temperature regimes for 3 months, simulating the temperature of the lake at 4 m and 15 m depth. Possible effects of food quality were tested at each temperature using four diets differing in fatty acid composition. Temperature played an important role as a trigger for spawning, and the type of diet clearly affected the reproductive investment. When the heterokont chromophyte alga Nannochloropsis limnetica, which is rich in polyunsaturated fatty acids (PUFAs) and long-chained PUFAs (>C18), was fed to mussels, an increased egg mass was obtained. This result was in contrast to that found when the green alga Scenedesmus obliquus and the cyanobacterium Aphanothece sp., both of which are deficient in long-chained PUFAs, were offered as food to the mussels. Such a PUFA-dependent food quality may affect reproduction in lakes. Food quality effects vary seasonally in a lake and may be most important in summer, when low-food- quality green algae and cyanobacteria are abundant. The low biochemical quality of these blooms may affect at least the later period of gametogenesis of D. polymorpha, which reproduces from June to August.
Settlement on suitable substrata under favourable environmental conditions is an important factor for a successful recruitment of adult populations of Dreissena polymorpha. Therefore, the pattern of settlement of zebra mussel (Dreissena polymorpha) larvae at different depths was studied in Lake Constance. Maximum densities of larvae and newly settled juvenile mussels were observed at 4m depth, while only single settlement episodes were recorded at greater depths (15 m and 30 m). Temperature fluctuation was used as a surrogate parameter for internal seiches. Biotic and abiotic parameters were subjected to a principal component analysis (PCA). The tight coupling of the internal seiches, larval abundance and settlement at 4-m depth vs. the lack of coupling of the latter two variables at greater depths indicated that water currents transported settling larvae to the substrata at greater depth. Our data suggest that physical factors, such as boundary mixing and internal seiches, should be considered as sources of variability in settlement.
Significant seasonal variation in size at settlement has been observed in newly settled larvae of Dreissena polymorpha in Lake Constance. Diet quality, which varies temporally and spatially in freshwater habitats, has been suggested as a significant factor influencing life history and development of freshwater invertebrates. Accordingly, experiments were conducted with field-collected larvae to test the hypothesis that diet quality can determine planktonic larval growth rates, size at settlement and subsequent post-metamorphic growth rates. Larvae were fed one of two diets or starved. One diet was composed of cyanobacterial cells which are deficient in polyunsaturated fatty acids (PUFAs), and the other was a mixed diet rich in PUFAs. Freshly metamorphosed animals from the starvation treatment had a carbon content per individual 70% lower than that of larvae fed the mixed diet. This apparent exhaustion of larval internal reserves resulted in a 50% reduction of the postmetamorphic growth rates. Growth was also reduced in animals previously fed the cyanobacterial diet. Hence, low food quantity or low food quality during the larval stage of D. polymorpha lead to irreversible effects for postmetamorphic animals, and is related to inferior competitive abilities.
Food quality effects of unsaturated fatty acids on larvae of the zebra mussel Dreissena polymorpha
(2002)
In standardized growth experiments, newly hatched larvae of the zebra mussel Dreissena polymorpha were fed diets representing different biochemical compositions. Algae that were rich in (n-3) polyunsaturated fatty acids (PUFAs), except for long-chained (.C18) PUFAs (Chlorella minutissima and Monoraphidium minutum) were of low food quality. Higher growth than on C. minutissima or M. minutum was supported by a culture of the cyanobacterium Aphanothece sp., which contained traces of a long-chained (n-3) PUFA, docosahexaenoic acid (DHA, 22 : 6n-3). The alga Isochrysis aff. galbana, which contained high amounts of the longchained (n-3) PUFAs DHA and eicosapentaenoic acid (EPA, 20 : 5n- 3), supported the highest growth. The alga Nannochloropsis limnetica, which differed from I. galbana by a defi- ciency in DHA, allowed slightly, but significantly lower, growth. Growth of larvae on N. limnetica was increased by enrichment of N. limnetica cells with a lipid extract of I. galbana, showing that larval growth on N. limnetica was limited by the deficiency of a compound that was present in I. galbana. Growth was also enhanced by feeding N. limnetica cells supplemented with DHA, but not by cells enriched with EPA, indicating that DHA was the limiting factor. We conclude that, on DHA-deficient food, the larvae of D. polymorpha were not able to sufficiently convert C18-PUFAs into long- chained (n-3) PUFAs and that the rates for elongation and desaturation of EPA into DHA limited growth.
A significant seasonal variation in size at settlement has been observed in newly settled larvae of Dreissena polymorpha in Lake Constance. Diet quality, which varies temporally and spatially in freshwater habitats, has been suggested as a significant factor influencing the life history and development of freshwater invertebrates. Accordingly, experiments were conducted with field-collected larvae to test the proposal that diet quality can determine planktonic larval growth rates, size at settlement and subsequent post-metamorphic growth rates. Larvae were fed one of two diets or starved. One diet was composed of cyanobacterial cells, which are de; cient in polyunsaturated fatty acids (PUFAs) and the other was a mixed diet rich in PUFAs. Freshly metamorphosed animals from the starvation treatment had a carbon content per individual 70% lower than that of larvae fed the mixed diet. This apparent exhaustion of larval internal reserves resulted in a 50% reduction of the post-metamorphic growth rates. Growth was also reduced in animals previously fed the cyanobacterial diet. Hence, low food quantity or low food quality during the larval stage of D. polymorpha, lead to irreversible effects for post-metamorphic animals and are related to inferior competitive abilities.
The factors that determine the efficiency of energy transfer in aquatic food webs have been investigated for many decades. The plant-animal interface is the most variable and least predictable of all levels in the food web. In order to study determinants of food quality in a large lake and to test the recently proposed central importance of the long-chained eicosapentaenoic acid (EPA) at the pelagic producer-grazer interface, we tested the importance of polyunsaturated fatty acids (PUFAs) at the pelagic producerconsumer interface by correlating sestonic food parameters with somatic growth rates of a clone of Daphnia galeata. Daphnia growth rates were obtained from standardized laboratory experiments spanning one season with Daphnia feeding on natural seston from Lake Constance, a large pre-alpine lake. Somatic growth rates were fitted to sestonic parameters by using a saturation function. A moderate amount of variation was explained when the model included the elemental parameters carbon (r2 = 0.6) and nitrogen (r2 = 0.71). A tighter fit was obtained when sestonic phosphorus was incorporated (r2 = 0.86). The nonlinear regression with EPA was relatively weak (r2 = 0.77), whereas the highest degree of variance was explained by three C18-PUFAs. The best (r2 = 0.95), and only significant, correlation of Daphnia's growth was found with the C18-PUFA α-linolenic acid (α-LA; C18:3n-3). This correlation was weakest in late August when C:P values increased to 300, suggesting that mineral and PUFA-limitation of Daphnia's growth changed seasonally. Sestonic phosphorus and some PUFAs showed not only tight correlations with growth, but also with sestonic α-LA content. We computed Monte Carlo simulations to test whether the observed effects of α-LA on growth could be accounted for by EPA, phosphorus, or one of the two C18-PUFAs, stearidonic acid (C18:4n-3) and linoleic acid (C18:2n-6). With >99 % probability, the correlation of growth with α-LA could not be explained by any of these parameters. In order to test for EPA limitation of Daphnia's growth, in parallel with experiments on pure seston, growth was determined on seston supplemented with chemostat-grown, P-limited Stephanodiscus hantzschii, which is rich in EPA. Although supplementation increased the EPA content 80-800x, no significant changes in the nonlinear regression of the growth rates with α-LA were found, indicating that growth of Daphnia on pure seston was not EPA limited. This indicates that the two fatty acids, EPA and α-LA, were not mutually substitutable biochemical resources and points to different physiological functions of these two PUFAs. These results support the PUFA-limitation hypothesis for sestonic C:P < 300 but are contrary to the hypothesis of a general importance of EPA, since no evidence for EPA limitation was found. It is suggested that the resource ratios of EPA and α-LA rather than the absolute concentrations determine which of the two resources is limiting growth.
The factors that determine the efficiency of energy transfer in aquatic food webs have been investigated for many decades. The plant-animal interface is the most variable and least predictable of all levels in the food web. In order to study determinants of food quality in a large lake and to test the recently proposed central importance of the long-chained eicosapentaenoic acid (EPA) at the pelagic producer-grazer interface, we tested the importance of polyunsaturated fatty acids (PUFAs) at the pelagic producer-consumer interface by correlating sestonic food parameters with somatic growth rates of a clone of Daphnia galeata. Daphnia growth rates were obtained from standardized laboratory experiments spanning one season with Daphnia feeding on natural seston from Lake Constance, a large pre-alpine lake. Somatic growth rates were fitted to sestonic parameters by using a saturation function. A moderate amount of variation was explained when the model included the elemental parameters carbon (r2 = 0.6) and nitrogen (r2 = 0.71). A tighter fit was obtained when sestonic phosphorus was incorporated (r2 = 0.86). The nonlinear regression with EPA was relatively weak (r2 = 0.77), whereas the highest degree of variance was explained by three C18-PUFAs. The best (r2 = 0.95), and only significant, correlation of Daphnia's growth was found with the C18-PUFA a-linolenic acid (a-LA; C18:3n-3). This correlation was weakest in late August when C:P values increased to 300, suggesting that mineral and PUFA- limitation of Daphnia's growth changed seasonally. Sestonic phosphorus and some PUFAs showed not only tight correlations with growth, but also with sestonic alpha-LA content. We computed Monte Carlo simulations to test whether the observed effects of alpha-LA on growth could be accounted for by EPA, phosphorus, or one of the two C18-PUFAs, stearidonic acid (C18:4n-3) and linoleic acid (C18:2n-6). With >99% probability, the correlation of growth with alpha-LA could not be explained by any of these parameters. In order to test for EPA limitation of Daphnia's growth, in parallel with experiments on pure seston, growth was determined on seston supplemented with chemostat-grown, Plimited Stephanodiscus hantzschii, which is rich in EPA. Although supplementation increased the EPA content 80-800x, no significant changes in the nonlinear regression of the growth rates with alpha-LA were found, indicating that growth of Daphnia on pure seston was not EPA limited. This indicates that the two fatty acids, EPA and alpha-LA, were not mutually substitutable biochemical resources and points to different physiological functions of these two PUFAs. These results support the PUFA-limitation hypothesis for sestonic C:P < 300 but are contrary to the hypothesis of a general importance of EPA, since no evidence for EPA limitation was found. It is suggested that the resource ratios of EPA and alpha-LA rather than the absolute concentrations determine which of the two resources is limiting growth.