Erik Sperfeld

Colimitation of a freshwater herbivore by sterols and polyunsaturated fatty acids

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.

Productivity in the Barents Sea - Response to Recent Climate Variability

The temporal and spatial dynamics of primary and secondary biomass/production in the Barents Sea since the late 1990s are examined using remote sensing data, observations and a coupled physical-biological model. Field observations of mesozooplankton biomass, and chlorophyll a data from transects (different seasons) and large-scale surveys (autumn) were used for validation of the remote sensing products and modeling results. The validation showed that satellite data are well suited to study temporal and spatial dynamics of chlorophyll a in the Barents Sea and that the model is an essential tool for secondary production estimates. Temperature, open water area, chlorophyll a, and zooplankton biomass show large interannual variations in the Barents Sea. The climatic variability is strongest in the northern and eastern parts. The moderate increase in net primary production evident in this study is likely an ecosystem response to changes in climate during the same period. Increased open water area and duration of open water season, which are related to elevated temperatures, appear to be the key drivers of the changes in annual net primary production that has occurred in the northern and eastern areas of this ecosystem. The temporal and spatial variability in zooplankton biomass appears to be controlled largely by predation pressure. In the southeastern Barents Sea, statistically significant linkages were observed between chlorophyll a and zooplankton biomass, as well as between net primary production and fish biomass, indicating bottom-up trophic interactions in this region.

Effects of temperature and dietary sterol availability on growth and cholesterol allocation of the aquatic keystone species Daphnia.

SUMMARY 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.

Bridging factorial and gradient concepts of resource co-limitation: towards a general framework applied to consumers.

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.

Bridging Ecological Stoichiometry and Nutritional Geometry with homeostasis concepts and integrative models of organism nutrition

Summary n1.The role of nutrition in linking animals with their environment is increasingly seen as fundamental to explain ecological interactions. n n2.The two currently predominant frameworks for exploring questions in nutritional ecology—Nutritional Geometry (NG) and Ecological Stoichiometry (ES)—share common features, but also differ in their goals and origins. NG originates from behavioural ecology using terrestrial insects as model organisms in tightly controlled feeding experiments, while ES originates from biogeochemistry focusing on the transfer of key elements across trophic levels, mainly in aquatic environments. n n3.Here, we review the history of these two complementary frameworks, emphasizing the key concepts defining their respective aims, methodologies, and focal taxa to answer questions at different ecological scales. n n4.We identify and explore homeostasis as a shared conceptual cornerstone of each framework that can be used to bridge knowledge gaps and for developing new hypotheses within nutritional ecology. n n5.Expanding on the concept of homeostasis, we introduce dynamic energy budget (DEB) models as a general way to address homeostatic regulation at its fundamental level. n n6.Specifically, we describe how a two-reserve DEB model can be used to track metabolic pathways of nutrients as well as elements and suggest that multi-reserve DEB models, when integrated and parameterized with NG and ES concepts, can form powerful components of agent-based models to predict how animal nutrition influences individual and trophic interactions in food webs. n nThis article is protected by copyright. All rights reserved.

Quantity and quality limit detritivore growth: mechanisms revealed by ecological stoichiometry and co-limitation theory

Resource quantity and quality are fundamental bottom-up constraints on consumers. Best understood in autotroph-based systems, co-occurrence of these constraints may be common but remains poorly studied in detrital-based systems. Here, we used a laboratory growth experiment to test limitation of the detritivorous caddisfly larvae Pycnopsyche lepida across a concurrent gradient of oak litter quantity (food supply) and quality (phosphorusu200a:u200acarbon [P:C ratios]). Growth increased simultaneously with quantity and quality, indicating co-limitation across the resource gradients. We merged approaches of ecological stoichiometry and co-limitation theory, showing how co-limitation reflected shifts in C and P acquisition throughout homeostatic regulation. Increased growth was best explained by elevated consumption rates and improved P assimilation, which both increased with elevated quantity and quality. Notably, C assimilation efficiencies remained unchanged and achieved maximum 18% at low quantity despite pronounced C limitation. Detrital C recalcitrance and substantive post-assimilatory C losses probably set a minimum quantity threshold to achieve positive C balance. Above this threshold, greater quality enhanced larval growth probably by improving P assimilation toward P-intensive growth. We suggest this interplay of C and P acquisition contributes to detritivore co-limitation, highlighting quantity and quality as potential simultaneous bottom-up controls in detrital-based ecosystems, including under anthropogenic change like nutrient enrichment.

Maternal diet of Daphnia magna affects offspring growth responses to supplementation with particular polyunsaturated fatty acids

Previous studies examining the effects of food quality on zooplankton often controlled for maternal effects of resource provisioning using standardized maternal diets. However, varying nutritional history of mothers may change resource provisioning to their progeny, especially regarding polyunsaturated fatty acids (PUFAs), which may change the interpretation of previously observed fitness responses of offspring. To assess PUFA-mediated maternal provisioning effects on offspring, we raised females of the cladoceran Daphnia magna on diets differing considerably in PUFA composition and raised their offspring on a PUFA-lacking diet supplemented with the ω3 PUFAs α-linolenic acid (ALA) and/or eicosapentaenoic acid (EPA). The mass-specific growth responses of offspring to their own diets were affected by the maternal diet regime, probably due to varying maternal PUFA provisioning. A low maternal provisioning of EPA or ALA was sufficient to prevent growth limitation of offspring by these PUFAs until reaching maturity. A comparison with results of published ALA and EPA supplementation experiments suggests that the previously observed limitation effects depended on the usage of a single algae genus as maternal diet. Therefore, we suggest that maternal diets should be deliberately varied in future studies assessing ecological relevant food quality effects on zooplankton, especially regarding PUFAs.

Predator–prey interactions in a changing world: humic stress disrupts predator threat evasion in copepods

Increasing inputs of colored dissolved organic matter (cDOM), which is mainly composed of humic substances (HS), are a widespread phenomenon of environmental change in aquatic ecosystems. This process of brownification alters the chemical conditions of the environment, but knowledge is lacking of whether elevated cDOM and HS levels interfere with the ability of prey species to evade chemical predator cues and thus affect predator–prey interactions. We assessed the effects of acute and prolonged exposure to HS at increasing concentrations on the ability of freshwater zooplankton to avoid predator threat (imposed by fish kairomones) in laboratory trials with two calanoid copepods (Eudiaptomus gracilis and Heterocope appendiculata). Populations of both species clearly avoided water containing fish kairomones. However, the avoidance behavior weakened with increasing HS concentration, suggesting that HS affected the ability of copepods to perceive or respond to the predator cue. The behavioral responses of the two copepod populations to increasing HS concentrations differed, with H. appendiculata being more sensitive than E. gracilis in an acute exposure scenario, whereas E. gracilis responded more strongly after prolonged exposure. Both showed similar physiological impairment after prolonged exposure, as revealed by their oxidative balance as a stress indicator, but mortality increased more strongly for H. appendiculata when the HS concentration increased. These results indicate that reduced predator threat evasion in the presence of cDOM could make copepods more susceptible to predation in future, with variation in the strength of responses among populations leading to changes in zooplankton communities and lake food-web structure.