@misc{WackervonElert2001,
  author    = {Wacker, Alexander and von Elert, Eric},
  title     = {Polyunsaturated fatty acids : evidence for non-substitutable biochemical resources in Daphnia galeata},
  url       = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus-17587},
  year      = {2001},
  abstract  = {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.},
  language  = {en}
}
@misc{WackervonElert2002,
  author    = {Wacker, Alexander and von Elert, Eric},
  title     = {Strong influences of larval diet history on subsequent post-settlement growth in the freshwater mollusc Dreissena polymorpha},
  url       = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus-17627},
  year      = {2002},
  abstract  = {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.},
  language  = {en}
}
@article{WackerMarzetzSpijkerman2015,
  author    = {Wacker, Alexander and Marzetz, Vanessa and Spijkerman, Elly},
  title     = {Interspecific competition in phytoplankton drives the availability of essential mineral and biochemical nutrients},
  series = {Ecology : a publication of the Ecological Society of America},
  volume    = {96},
  journal   = {Ecology : a publication of the Ecological Society of America},
  number    = {9},
  publisher = {Wiley},
  address   = {Washington},
  issn      = {0012-9658},
  doi       = {10.1890/14-1915.1},
  pages     = {2467 -- 2477},
  year      = {2015},
  abstract  = {The underlying mechanisms and consequences of competition and diversity are central themes in ecology. A higher diversity of primary producers often results in higher resource use efficiency in aquatic and terrestrial ecosystems. This may result in more food for consumers on one hand, while, on the other hand, it can also result in a decreased food quality for consumers; higher biomass combined with the same availability of the limiting compound directly reduces the dietary proportion of the limiting compound. Here we tested whether and how interspecific competition in phytoplankton communities leads to changes in resource use efficiency and cellular concentrations of nutrients and fatty acids. The measured particulate carbon : phosphorus ratios (C:P) and fatty acid concentrations in the communities were compared to the theoretically expected ratios and concentrations of measurements on simultaneously running monocultures. With interspecific competition, phytoplankton communities had higher concentrations of the monounsaturated fatty acid oleic acid and also much higher concentrations of the ecologically and physiologically relevant long-chain polyunsaturated fatty acid eicosapentaenoic acid than expected concentrations based on monocultures. Such higher availability of essential fatty acids may contribute to the positive relationship between phytoplankton diversity and zooplankton growth, and may compensate limitations by mineral nutrients in higher trophic levels.},
  language  = {en}
}
@article{SperfeldWacker2012,
  author    = {Sperfeld, Erik and Wacker, Alexander},
  title     = {Temperature affects the limitation of Daphnia magna by eicosapentaenoic acid, and the fatty acid composition of body tissue and eggs},
  series = {Freshwater biology},
  volume    = {57},
  journal   = {Freshwater biology},
  number    = {3},
  publisher = {Wiley-Blackwell},
  address   = {Hoboken},
  issn      = {0046-5070},
  doi       = {10.1111/j.1365-2427.2011.02719.x},
  pages     = {497 -- 508},
  year      = {2012},
  abstract  = {1. Poikilothermic animals incorporate more polyunsaturated fatty acids (PUFAs) into their cellular membranes as temperature declines, suggesting an increased sensitivity to PUFA limitation in cool conditions. To test this we raised Daphnia magna at different temperatures and investigated the effect of varying dietary PUFA on life history parameters (i.e. growth, reproduction) and the PUFA composition of body tissue and eggs. 2. Upon a PUFA-rich diet (Cryptomonas sp.) females showed higher concentrations of several omega 3 PUFAs in their body tissue at 15 degrees C than at 20 degrees C and 25 degrees C, indicating a greater structural requirement for omega 3 PUFAs at low temperature. Their eggs had an equal but higher concentration of omega 3 PUFAs than their body tissue. 3. In a life history experiment at 15 and 20 degrees C we supplemented a diet of a PUFA-free cyanobacterium with the omega 3 PUFA eicosapentaenoic acid (EPA). The growth of D. magna was more strongly EPA limited at low temperature. A greater requirement for structural EPA at 15 degrees C was indicated by a steeper increase in somatic EPA content with dietary EPA compared to 20 degrees C. 4. At 20 degrees C the development of eggs to successful hatching was high when EPA was supplied to the mothers. At 15 degrees C the hatching success was generally poor, despite of a higher maternal provision of EPA to eggs, compared to that at 20 degrees C, suggesting that EPA alone was insufficient for proper neonatal development at the low temperature. The growth of offspring from mothers raised at 20 degrees C without EPA supplementation was very low, indicating that the negative effects of EPA deficiency can be carried on to the next generation. 5. The fatty acid composition of Daphnia sp. in published field studies shows increasing proportions of saturated fatty acids with increasing environmental temperature, whereas omega 3 PUFAs and EPA show no clear pattern, suggesting that variations in dietary PUFA may mask temperature-dependent adjustments in omega 3 PUFA concentrations of cladocerans in nature.},
  language  = {en}
}
@misc{SperfeldRaubenheimerWacker2016,
  author    = {Sperfeld, Erik and Raubenheimer, David and Wacker, Alexander},
  title     = {Bridging factorial and gradient concepts of resource co-limitation: towards a general framework applied to consumers},
  series = {Ecology letters},
  volume    = {19},
  journal   = {Ecology letters},
  publisher = {Wiley-Blackwell},
  address   = {Hoboken},
  issn      = {1461-023X},
  doi       = {10.1111/ele.12554},
  pages     = {201 -- 215},
  year      = {2016},
  abstract  = {Organism growth can be limited either by a single resource or by multiple resources simultaneously (co-limitation). Efforts to characterise co-limitation have generated two influential approaches. One approach uses limitation scenarios of factorial growth assays to distinguish specific types of co-limitation; the other uses growth responses spanned over a continuous, multi-dimensional resource space to characterise different types of response surfaces. Both approaches have been useful in investigating particular aspects of co-limitation, but a synthesis is needed to stimulate development of this recent research area. We address this gap by integrating the two approaches, thereby presenting a more general framework of co-limitation. We found that various factorial (co-)limitation scenarios can emerge in different response surface types based on continuous availabilities of essential or substitutable resources. We tested our conceptual co-limitation framework on data sets of published and unpublished studies examining the limitation of two herbivorous consumers in a two-dimensional resource space. The experimental data corroborate the predictions, suggesting a general applicability of our co-limitation framework to generalist consumers and potentially also to other organisms. The presented framework might give insight into mechanisms that underlie co-limitation responses and thus can be a seminal starting point for evaluating co-limitation patterns in experiments and nature.},
  language  = {en}
}
@article{SperfeldMartinCreuzburgWacker2012,
  author    = {Sperfeld, Erik and Martin-Creuzburg, Dominik and Wacker, Alexander},
  title     = {Multiple resource limitation theory applied to herbivorous consumers Liebig's minimum rule vs. interactive co-limitation},
  series = {Ecology letters},
  volume    = {15},
  journal   = {Ecology letters},
  number    = {2},
  publisher = {Wiley-Blackwell},
  address   = {Malden},
  issn      = {1461-023X},
  doi       = {10.1111/j.1461-0248.2011.01719.x},
  pages     = {142 -- 150},
  year      = {2012},
  abstract  = {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.},
  language  = {en}
}
@article{SchaelickeHeimMartinCreuzburgetal.2020,
  author    = {Sch{\"a}licke, Svenja and Heim, Silvia and Martin-Creuzburg, Dominik and Wacker, Alexander},
  title     = {Inter- and intraspecific differences in rotifer fatty acid composition during acclimation to low-quality food},
  series = {Philosophical transactions of the Royal Society of London : B, Biological sciences},
  volume    = {375},
  journal   = {Philosophical transactions of the Royal Society of London : B, Biological sciences},
  number    = {1804},
  publisher = {Royal Society},
  address   = {London},
  issn      = {0962-8436},
  doi       = {10.1098/rstb.2019.0644},
  pages     = {8},
  year      = {2020},
  abstract  = {Biochemical food quality constraints affect the performance of consumers and mediate trait variation among and within consumer species. To assess inter- and intraspecific differences in fatty acid retention and conversion in freshwater rotifers, we provided four strains of two closely related rotifer species,Brachionus calyciflorussensustricto andBrachionus fernandoi, with food algae differing in their fatty acid composition. The rotifers grazed for 5 days on eitherNannochloropsis limneticaorMonoraphidium minutum, two food algae with distinct polyunsaturated fatty acid (PUFA) profiles, before the diets were switched to PUFA-freeSynechococcus elongatus, which was provided for three more days. We found between- and within-species differences in rotifer fatty acid compositions on the respective food sources and, in particular, highly specific acclimation reactions to the PUFA-free diet. The different reactions indicate inter- but also intraspecific differences in physiological traits, such as PUFA retention, allocation and bioconversion capacities, within the genusBrachionusthat are most likely accompanied by differences in their nutritional demands. Our data suggest that biochemical food quality constraints act differently on traits of closely related species and of strains of a particular species and thus might be involved in shaping ecological interactions and evolutionary processes. This article is part of the theme issue 'The next horizons for lipids as 'trophic biomarkers': evidence and significance of consumer modification of dietary fatty acids'.},
  language  = {en}
}
@misc{MartinCreuzburgMassierWacker2018,
  author    = {Martin-Creuzburg, Dominik and Massier, Tamara and Wacker, Alexander},
  title     = {Sex-specific differences in essential lipid requirements of Daphnia magna},
  series = {Postprints der Universit{\"a}t Potsdam : Mathematisch-Naturwissenschaftliche Reihe},
  journal   = {Postprints der Universit{\"a}t Potsdam : Mathematisch-Naturwissenschaftliche Reihe},
  number    = {1050},
  issn      = {1866-8372},
  doi       = {10.25932/publishup-46909},
  url       = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus4-469099},
  pages     = {16},
  year      = {2018},
  abstract  = {Sex-specific differences in nutritional requirements may crucially influence the performances of the sexes, which may have implications for sexual reproduction and thus is of great ecological and evolutionary interest. In the freshwater model species Daphnia magna, essential lipid requirements have been extensively studied. Dietary deficiencies in sterols and polyunsaturated fatty acids (PUFA) have been shown to constrain somatic growth and parthenogenetic reproduction of female Daphnia. In contrast, nutrient requirements of male Daphnia have not been studied yet. Supplementation experiments were conducted to investigate differences in sterol (cholesterol) and PUFA (eicosapentaenoic acid, EPA) requirements between female and male D. magna. Thresholds for sterol-limited juvenile growth were higher in females than in males, suggesting that females are more susceptible to dietary sterol deficiencies than males. Sex-specific differences in maximum somatic growth rates were evident primarily in the presence of dietary EPA; females could not exploit their generally higher growth potential in the absence of dietary PUFA. However, the thresholds for EPA-limited growth did not differ between sexes, suggesting that both sexes have similar dietary EPA requirements during juvenile growth. During a life history experiment, the gain in body dry mass was higher in females than in males, irrespective of food treatment. In both sexes, the gain in body dry mass increased significantly upon EPA supplementation, indicating that both sexes benefited from dietary EPA supply also later in life. However, the positive effects of EPA supplementation were most pronounced for female reproduction-related traits (i.e., clutch sizes, egg dry masses, and total dry mass investment in reproduction). The high maternal investment in reproduction resulted in a depletion of nutrients in female somata. In contrast, the comparatively low paternal investment in reproduction allowed for the accumulation of nutrients in male somata. We conclude that males are generally less susceptible to dietary nutrient deficiencies than females, because they can rely more on internal body stores. Our data suggest that the performances of the sexes are differentially influenced by lipid-mediated food quality, which may have consequences for sexual reproduction and thus the production of resting eggs and the maintenance of Daphnia populations.},
  language  = {en}
}
@article{MartinCreuzburgMassierWacker2018,
  author    = {Martin-Creuzburg, Dominik and Massier, Tamara and Wacker, Alexander},
  title     = {Sex-Specific differences in essential lipid requirements of Daphnia magna},
  series = {Frontiers in Ecology and Evolution},
  volume    = {6},
  journal   = {Frontiers in Ecology and Evolution},
  publisher = {Frontiers Research Foundation},
  address   = {Lausanne},
  issn      = {2296-701X},
  doi       = {10.3389/fevo.2018.00089},
  pages     = {14},
  year      = {2018},
  abstract  = {Sex-specific differences in nutritional requirements may crucially influence the performances of the sexes, which may have implications for sexual reproduction and thus is of great ecological and evolutionary interest. In the freshwater model species Daphnia magna, essential lipid requirements have been extensively studied. Dietary deficiencies in sterols and polyunsaturated fatty acids (PUFA) have been shown to constrain somatic growth and parthenogenetic reproduction of female Daphnia. In contrast, nutrient requirements of male Daphnia have not been studied yet. Supplementation experiments were conducted to investigate differences in sterol (cholesterol) and PUFA (eicosapentaenoic acid, EPA) requirements between female and male D. magna. Thresholds for sterol-limited juvenile growth were higher in females than in males, suggesting that females are more susceptible to dietary sterol deficiencies than males. Sex-specific differences in maximum somatic growth rates were evident primarily in the presence of dietary EPA; females could not exploit their generally higher growth potential in the absence of dietary PUFA. However, the thresholds for EPA-limited growth did not differ between sexes, suggesting that both sexes have similar dietary EPA requirements during juvenile growth. During a life history experiment, the gain in body dry mass was higher in females than in males, irrespective of food treatment. In both sexes, the gain in body dry mass increased significantly upon EPA supplementation, indicating that both sexes benefited from dietary EPA supply also later in life. However, the positive effects of EPA supplementation were most pronounced for female reproduction-related traits (i.e., clutch sizes, egg dry masses, and total dry mass investment in reproduction). The high maternal investment in reproduction resulted in a depletion of nutrients in female somata. In contrast, the comparatively low paternal investment in reproduction allowed for the accumulation of nutrients in male somata. We conclude that males are generally less susceptible to dietary nutrient deficiencies than females, because they can rely more on internal body stores. Our data suggest that the performances of the sexes are differentially influenced by lipid-mediated food quality, which may have consequences for sexual reproduction and thus the production of resting eggs and the maintenance of Daphnia populations.},
  language  = {en}
}
@article{LukasWacker2014,
  author    = {Lukas, Marcus and Wacker, Alexander},
  title     = {Acclimation to dietary shifts impacts the carbon budgets of Daphnia magna},
  series = {Journal of plankton research},
  volume    = {36},
  journal   = {Journal of plankton research},
  number    = {3},
  publisher = {Oxford Univ. Press},
  address   = {Oxford},
  issn      = {0142-7873},
  doi       = {10.1093/plankt/fbu018},
  pages     = {848 -- 858},
  year      = {2014},
  abstract  = {Daphnia responds to low availability of carbon (food quantity) or limiting concentrations of nutrients relative to carbon (C) in excess (food quality) by respectively saving or discharging C via different pathways. We investigated which kind of food limitation leads to a faster regulation in Daphnia C budgets, and whether the pre-assimilative C pathways, ingestion and faeces egestion and the post-assimilative C pathways, excretion and respiration, are regulated concurrently. Daphnia magna were exposed to dietary shifts in different food quantities or qualities; food quality was varied in terms of the essential component, cholesterol. After acclimation to the new diet ranging from 0 to 96 h, C budgets were measured by a radiotracer technique. Dietary shifts in quantity and quality caused Daphnia to quickly adjust their C budgets within 6 h, but different C pathways were affected. A shift to low food quantity reduced Daphnia respiration indicating C retention. In contrast, sudden low quality food caused increased faeces egestion to discharge excess C. Furthermore, we observed a delayed increase in excretion but no change in respiration within the time frame studied. Such time-shifted responses appear to be an appropriate means to keep the costs of physiological adjustments relatively low, which in turn would benefit Daphnia performance.},
  language  = {en}
}