Food quantity-quality interactions determine growth rates and reproductive success of consumers and thereby regulate community dynamics and food web structure. Predator-prey models that shape our conceptual understanding of foraging ecology typically rely on the parametrization of fixed consumer responses to either food quantity or food quality. In nature, however, consumers optimize their fitness by responding simultaneously to changes in food quantity and quality. Therefore, we assessed consumer responses to changing food environments using a new fitness optimization model that accounted for food quality-quantity interactions to better capture the regulatory flexibility of consumers. Our simulations demonstrated that the impact of food quality on important consumer traits can be altered or even reversed by changes in food quality. Low food quality, for example, affected feeding rates negatively at low food concentrations but triggered surplus feeding at high food concentrations. The scope of surplus feeding was thereby mainly dependent on dynamics of nutrient digestion and in contrast to previous assumptions, energy costs of feeding played a minor role. Further, the regulation of digestive enzyme production, a crucial factor determining assimilation efficiencies, was strongly dependent on whether nonessential or essential nutrients were limiting growth. Consequently, not only the degree but also the type of nutrient limitation mediated the impact of the food environment on consumers' fitness. At the community level, food quality was key in shaping predator-prey biomass ratios. High food qualities resulted in top-heavy systems with larger consumer than prey biomass. Decreases of prey digestibility or the availability of essential nutrients, however, triggered a switch from inverted to classical pyramid shapes of bi-trophic systems. The impact of food quantity on trophic transfer and emerging structural ecosystem properties thus critically hinges on behavioral and physiological responses of consumers. The inclusion of the regulatory flexibility of consumers is therefore an essential next step to improve predator-prey models and our conceptual understanding of trophic interactions.

Compound-specific isotope analyses (CSIA) of fatty acids (FA) constitute a promising tool for tracing energy flows in food-webs. However, past applications of FA-specific carbon isotope analyses have been restricted to a relatively coarse food-source separation and mainly quantified dietary contributions from different habitats. Our aim was to evaluate the potential of FA-CSIA to provide high-resolution data on within-system energy flows using algae and zooplankton as model organisms. First, we investigated the power of FA-CSIA to distinguish among four different algae groups, namely cyanobacteria, chlorophytes, haptophytes and diatoms. We found substantial within-group variation but also demonstrated that delta C-13 of several FA (e.g. 18:3 omega 3 or 18:4 omega 3) differed among taxa, resulting in group-specific isotopic fingerprints. Second, we assessed changes in FA isotope ratios with trophic transfer. Isotope fractionation was highly variable in daphnids and rotifers exposed to different food sources. Only delta C-13 of nutritionally valuable poly-unsaturated FA remained relatively constant, highlighting their potential as dietary tracers. The variability in fractionation was partly driven by the identity of food sources. Such systematic effects likely reflect the impact of dietary quality on consumers' metabolism and suggest that FA isotopes could be useful nutritional indicators in the field. Overall, our results reveal that the variability of FA isotope ratios provides a substantial challenge, but that FA-CSIA nevertheless have several promising applications in food-web ecology. 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'.

Estimating trophic structures is a common approach used to retrieve information regarding energy pathways, predation, and competition in complex ecosystems. The application of amino acid (AA) compound-specific nitrogen (N) isotope analysis (CSIA) is a relatively new method used to estimate trophic position (TP) and feeding relationships in diverse organisms. Here, we conducted the first meta-analysis of delta N-15 AA values from measurements of 359 marine species covering four trophic levels, and compared TP estimates from AA-CSIA to literature values derived from food items, gut or stomach content analysis. We tested whether the AA trophic enrichment factor (TEF), or the N-15 enrichment among different individual AAs is constant across trophic levels and whether inclusion of delta N-15 values from multiple AAs improves TP estimation. For the TEF of glutamic acid relative to phenylalanine (Phe) we found an average value of 6.6 aEuro degrees across all taxa, which is significantly lower than the commonly applied 7.6 aEuro degrees. We found that organism feeding ecology influences TEF values of several trophic AAs relative to Phe, with significantly higher TEF values for herbivores compared to omnivores and carnivores, while TEF values were also significantly lower for animals excreting urea compared to ammonium. Based on the comparison of multiple model structures using the metadata of delta N-15 AA values we show that increasing the number of AAs in principle improves precision in TP estimation. This meta-analysis clarifies the advantages and limitations of using individual delta N-15 AA values as tools in trophic ecology and provides a guideline for the future application of AA-CSIA to food web studies.

Reliable tracer techniques are fundamental to retrieving accurate information on consumer dietary resource use in dynamic ecosystems. Analysis of delta C-13 values in essential amino acids has shown great promise in effectively capturing consumer food sources, since essential amino acids are not synthesized by heterotrophic organisms but instead routed directly from dietary sources to consumers, resulting in negligible C-13 trophic discrimination. We assessed seasonal feeding patterns of a widespread key copepod species (Acartia spp.) in the northern Baltic proper using complementary approaches: bulk delta C-13 and delta N-15 values, delta C-13 values of essential amino acids, and quantitative phytoplankton taxonomic data. Our results showed distinct differences between Acartia and seston delta C-13 essential amino acid values measured at weekly to monthly sampling intervals, which indicated that Acartia preferentially utilized specific dietary resources that comprised only parts of the total phytoplankton composition (varying from 19.7% to 81.4% during the season). Results also indicated that care should be taken when inferring trophic position from bulk stable isotopes when consumers are highly selective, since isotope values of seston may not accurately reflect consumer specific diet resource uptake. Analysis of delta C-13 values in essential amino acids was shown to be a promising tool to accurately trace consumer resource use in complex natural systems.

Food webs are structured by intricate nodes of species interactions which govern the flow of organic matter in natural systems. Despite being long recognized as a key component in ecology, estimation of food web functioning is still challenging due to the difficulty in accurately measuring species interactions within a food web. Novel tracing methods that estimate species diet uptake and trophic position are therefore needed for assessing food web dynamics.

The focus of this thesis is the use of compound specific nitrogen and carbon stable isotopes and molecular techniques for assessing predator-prey interactions and energy flow in natural aquatic ecosystems, with a particular focus on the species links between phytoplankton and zooplankton.

The use of δ¹⁵N amino acid values to predict organism trophic position are evaluated through a meta-analysis of available literature which included measurements from 359 marine species (article I). Through a controlled feeding study isotope incorporation in aquatic organisms, across both plant-animal and animal-animal species linkages is further assessed (article II).

These studies showed that δ¹⁵N amino acid values are useful tools for categorizing animal trophic position. Organism feeding ecology influenced nitrogen trophic discrimination (difference in isotope ratio between consumer and diet), with higher discrimination in herbivores compared to omnivores and carnivores (article I). Nitrogen isotope trophic discrimination also varied among feeding treatments in the laboratory study (article II). The combined findings from articles I & II suggest that researchers should consider using group specific nitrogen trophic discrimination values to improve accuracy in species trophic position predictions. 

Another key finding in the controlled laboratory study (article II) was consistently low carbon isotope discrimination in essential amino acids across all species linkages, confirming that these compounds are reliable dietary tracers.

The δ¹³C ratios of essential amino acids were applied to study seasonal dynamics in zooplankton resource use in the Baltic Sea (article III). Data from this study indicated that zooplankton assimilate variable resources throughout the growing season. Molecular diet analysis (article IV) showed that marine copepod and cladoceran species ingested both autotrophic and heterotrophic resources.

Evidence from both articles III & IV also revealed that zooplankton feed on a relatively broad range of diet items but not opportunistically on all available food sources. Mesozooplankton feeding patterns suggested that energy and nutritional flows were channelled through an omnivorous zooplankton food web including microzooplankton prey items. Overall the results of this thesis highlight that stable isotope ratios in specific compounds and molecular techniques are useful tracing approaches that improve our understanding of food web functioning.

Data on optical properties such as diffuse attenuation coefficient K (d)(PAR), beam attenuation coefficient (c (p)) and the optically active constituents (OACs) CDOM, Chl-a and suspended particulate matter were obtained in a Danish temperate coastal plain estuary (56A degrees N) and a Vietnamese tropical ria (12A degrees N) at high discharges. The major difference was the spatial distribution of the optical properties against distance, best described by significant power functions in the ria, compared to significant linear functions in the coastal plain. It was hypothesized that estuarine morphometry could explain this spatial distribution. Partition and multiple regression analyses showed that Chl-a governed K (d)(PAR) and beam attenuation coefficient in both estuaries. Significant, high correlations were obtained by multiple regression analyses in the estimation of K (d)(PAR) and beam attenuation coefficients in the two estuaries using OACs as input parameters. It is concluded that there are no large differences in OAC concentrations between the two estuaries. The spatial distributions of OACs and optical properties were significantly different and governed by the estuary morphometry, i.e. a power distribution in the tropical ria and a linear function in the temperate coastal plain estuary, and applicable to similar estuary types.