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Organisms at the base of stream food webs are typically poor in long‐chain polyunsaturated fatty acids (LC‐PUFA), especially in docosahexaenoic acid (DHA), whereas consumers at higher trophic levels are often rich in LC‐PUFA. For example, fish tissues, especially the brain, are DHA‐rich. This obvious mismatch between consumer LC‐PUFA and their basal dietary supply may result from selective retention and/or endogenous conversion of dietary precursors to LC‐PUFA.To determine which is more likely, we investigated compound‐specific carbon stable isotopes in PUFA (δ13CPUFA) of potential basal resources (stream epilithon, leaf litter) and consumers (invertebrates, European bullhead, and two salmonid species and their brain, eye, liver, and muscle tissues). We predicted that consumer‐PUFA, depleted in 13C values relative to their dietary sources, would indicate internal de novo PUFA synthesis. Alternatively, higher consumer‐δ13CPUFA values would imply selective retention of aquatic (epilithon and conditioned leaves) rather than terrestrial resources or internal production, irrespective of trophic levels.Invertebrate grazers and predators resembled δ13C values of essential fatty acids (δ13CEFA) of benthic algae, while shredder‐δ13CEFA values reflected those of conditioned rather than fresh leaves. Lower eye‐δ13CEFA values of salmonids than in livers indicated high retention of dietary PUFA sources (invertebrates and epilithon). Stable isotope values of eicosapentaenoic acid suggest that all consumers retained algal EPA, while insectivorous fish produced DHA in their liver. There is no further evidence from carbon stable isotopes for local PUFA conversion within neural tissues.Our study demonstrates that δ13CPUFA can be used to track sources of these highly functional molecules in aquatic consumers and highlights the importance of algal derived‐PUFA for these consumers in oligotrophic headwater streams. [ABSTRACT FROM AUTHOR]

Fatty acids (FA) in lake zooplankton are largely provided by dietary FA and affected by taxonomic composition of zooplankton, temperature, and other environmental conditions. However, the extent to which dietary FA are retained and/or bioconverted in zooplankton remains unclear. In this lake study, we examined how dietary FA (seston), temperature and physico‐chemical variables (i.e., concentrations of chlorophyll a, dissolved organic carbon, nitrate, ammonium, phosphate) explained the FA variation in membrane (phospholipids; PL) and storage (triacylglycerols; TAG) lipids of zooplankton.We tested the hypotheses that: (1) increasing dietary FA increases the FA content stored as TAG by zooplankton, whereas the FA content in PL remains unaffected by naturally occurring changes in dietary FA supply; and (2) FA in seston better predict the FA variations stored by zooplankton as TAG, whereas lake temperature better predicts the FA variations in zooplankton PL.Results show that an increase in dietary polyunsaturated FA increased polyunsaturated FA in TAG and PL of zooplankton, but not eicosapentaenoic acid or docosahexaenoic acid in PL. Moreover, the FA content of seston was more similar to that of zooplankton TAG than it was to the content of PL, indicating major taxon‐specific differences in membrane FA.Dietary FA are a major predictor of FA in TAG of zooplankton, while the taxonomic composition of zooplankton is a major predictor of FA in PL. Temperature and physico‐chemical variables explained little of the variations in zooplankton FA. This field study improves our understanding of how FA in PL and TAG of different zooplankton taxa respond to changes of dietary FA, temperature, concentrations of chlorophyll‐a, dissolved organic carbon, nitrate, ammonium, and phosphate to fulfil their physiological requirements. [ABSTRACT FROM AUTHOR]

We studied how physiologically important long‐chain polyunsaturated fatty acids (PUFA) in benthic macroinvertebrates (Asellus aquaticus, Chironomidae, and Oligochaeta) were related to those in periphyton and terrestrial organic matter (tree leaves), collected from littoral areas of 17 boreal lakes that differed in their dissolved organic carbon (DOC) and nutrient (phosphorus and nitrogen) concentrations. We also analyzed fatty acid (FA)‐specific stable carbon isotopes (δ13CFA) to investigate the dietary origin (periphyton vs. terrestrial organic matter) of PUFA in the consumers. In contrast to periphyton, terrestrial organic matter was deprived of long‐chain PUFA, such as eicosapentaenoic acid (EPA), but rich in short‐chain PUFA. The FA composition of macroinvertebrates was primarily taxon‐specific despite the large differences in DOC and nutrient concentrations of the lakes. An increase in DOC concentration had a negative impact on the EPA content of Asellus, chironomids, and oligochaetes as well as the total FA content of chironomids and oligochaetes. However, the FA content of macroinvertebrates was not related to lake total phosphorus concentrations, although the total FA and EPA content of periphyton increased with the trophic status of the study lakes. The δ13CPUFA values of macroinvertebrates were positively related with the δ13CPUFA of periphyton and weakly with δ13CPUFA of terrestrial leaf material. The results indicate that EPA in the studied macroinvertebrate taxa was mainly derived from an algal‐based diet and not via biosynthesis from allochthonous precursor FA. Thus, macroinvertebrate production in lakes may be limited by the available algae‐based food sources. [ABSTRACT FROM AUTHOR]

Consumers feeding at the aquatic–terrestrial ecosystem interface may obtain a mixture of aquatic and terrestrial diet resources that vary in nutritional composition. However, in lake riparian spiders, the relative significance of aquatic versus terrestrial diet sources remains to be explored. We investigated the trophic transfer of lipids and polyunsaturated fatty acids (PUFA) from emergent aquatic and terrestrial insects to spiders at varying distances from the shoreline of a subalpine lake in Austria, using differences in fatty acid profiles and compound‐specific stable carbon (δ13C) and hydrogen (δ2H) isotopes. The omega‐3 PUFA content of emergent aquatic insects was higher than that of terrestrial insects. Emergent aquatic insects contained on average 6.6 times more eicosapentaenoic acid (EPA) and 1.2 times more α‐linolenic acid (ALA) than terrestrial insects, whereas terrestrial insects contained on average 2.6 times more linoleic acid (LIN) than emergent aquatic insects. Spiders sampled directly on the lake and in upland habitats had similar EPA contents, but this EPA was derived from different diet sources, depending on the habitat. The δ13CEPA and δ2HEPA values of 'lake spiders' revealed an aquatic diet pathway (i.e. EPA of aquatic origin). In contrast, EPA of spiders collected in terrestrial habitats was depleted in both 13C and 2H compared to any potential food sources, and their ALA isotopic values, suggesting that EPA was partly bioconverted from its dietary precursor ALA (i.e. internal pathway). The δ2H values of fatty acids clearly indicated that diet sources differed depending on the spider's habitat, which was less evident from the δ13C values of the fatty acids. Our data highlight that spiders can use two distinct pathways (trophic versus metabolic) to satisfy their physiological EPA demand, depending on habitat use and dietary availability. [ABSTRACT FROM AUTHOR]

Organisms at the base of aquatic food webs synthesize essential nutrients, such as omega‐3 polyunsaturated fatty acids (n‐3 PUFA), which are transferred to consumers at higher trophic levels. Many consumers, requiring n‐3 long‐chain (LC) PUFA, such as eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA), have limited ability to biosynthesize them from the essential dietary precursor α‐linolenic acid (ALA) and thus rely on dietary provision of LC‐PUFA.We investigated LC‐PUFA metabolism in freshwater zooplankton using stable hydrogen isotopes (δ2H) of fatty acids as tracers. We conducted feeding experiments with the freshwater keystone grazer Daphnia to quantify changes in the δ2H value of body FA in response to the FA composition of their food and the δ2H value of the ambient water.The isotopic composition of LC‐PUFA changed in Daphnia, depending on the integration of 2H from ambient water during de novo synthesis or bioconversion from dietary precursors, allowing us to distinguish dietary from bioconverted EPA in body tissue. We tested the applicability of these laboratory findings in a field setting by analysing δ2H values of PUFA in primary producers and consumers in eutrophic ponds to track EPA sources of zooplankton.Multilinear regression models that included conversion of ALA to EPA correlated better with zooplankton δ2HEPA than seston δ2HEPA at low dietary EPA supply.This study provides evidence that zooplankton can compensate for low dietary EPA supply by activating LC‐PUFA biosynthesis and shows that herbivorous zooplankton play a crucial role in upgrading FA for higher trophic levels during low dietary EPA supply. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

In recent decades, much conceptual thinking in trophic ecology has been guided by theories of nutrient limitation and the flow of elements, such as carbon and nitrogen, within and among ecosystems. More recently, ecologists have also turned their attention to examining the value of specific dietary nutrients, in particular polyunsaturated fatty acids (PUFA), among which the omega‐3 PUFA, especially eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA) play a central role as essential components of neuronal cell membranes in many organisms. This review focuses on a new neuro‐ecological approach stemming from the biochemical (mechanistic) and physiological (functional) role of DHA in neuronal cell membranes, in particular in conjunction with G‐protein coupled receptors (GPCRs). We link the co‐evolution of these neurological functions to metabolic dependency on dietary omega‐3 PUFA. We outline ways in which deficiencies in dietary DHA supply may affect, cognition, vision, and behaviour, and ultimately, the biological fitness of consumers. We then review emerging evidence that changes in access to dietary omega‐3 PUFA may ultimately have profound impacts on trophic interactions leading to potential changes in community structure and ecosystem functioning that, in turn, may affect the supply of DHA within and across ecosystems, including the supply for human consumption. [ABSTRACT FROM AUTHOR]

The nutritional diversity of resources can affect the adaptive evolution of consumer metabolism and consumer diversification. The omega‐3 long‐chain polyunsaturated fatty acids eicosapentaenoic acid (EPA; 20:5n‐3) and docosahexaenoic acid (DHA; 22:6n‐3) have a high potential to affect consumer fitness, through their widespread effects on reproduction, growth and survival. However, few studies consider the evolution of fatty acid metabolism within an ecological context. In this review, we first document the extensive diversity in both primary producer and consumer fatty acid distributions amongst major ecosystems, between habitats and amongst species within habitats. We highlight some of the key nutritional contrasts that can shape behavioural and/or metabolic adaptation in consumers, discussing how consumers can evolve in response to the spatial, seasonal and community‐level variation of resource quality. We propose a hierarchical trait‐based approach for studying the evolution of consumers' metabolic networks and review the evolutionary genetic mechanisms underpinning consumer adaptation to EPA and DHA distributions. In doing so, we consider how the metabolic traits of consumers are hierarchically structured, from cell membrane function to maternal investment, and have strongly environment‐dependent expression. Finally, we conclude with an outlook on how studying the metabolic adaptation of consumers within the context of nutritional landscapes can open up new opportunities for understanding evolutionary diversification. [ABSTRACT FROM AUTHOR]

Algae are generally a high‐quality diet source because they provide essential compounds to aquatic consumers. In forested stream ecosystems, the availability of high‐quality algae is low compared to terrestrial organic matter, which may constrain the dietary transfer of essential compounds to consumers. However, there could be other overlooked high‐quality resource pools that provide essential compounds to consumers in river ecosystems.We conducted a field study along a subalpine river continuum in Austria to identify the nutritional role of a 'hidden' food resource for aquatic consumers; the biofilms growing on the underside of rocks (dark biofilms). Dark and light (i.e. upper surface of rocks) biofilms and invertebrates were collected, and their fatty acid (FA) composition was analysed.Compared with light biofilms, dark biofilms contained greater proportions of bacterial FA, long‐chain saturated FA (biomarkers of terrestrial plants) and oleic acid (18:1ω9; a fungal biomarker), but a lower proportion of algal FA, especially omega‐3 polyunsaturated FA (ω3 PUFA). The ω3 PUFA composition in dark biofilms was strongly correlated with that in light biofilms. Furthermore, the overall FA profiles of dark biofilms were significantly associated with invertebrate FA profiles. Strong correlations were also observed between invertebrates and dark biofilms for bacterial FA and the ω3 PUFA eicosapentaenoic acid (EPA, 20:5ω3).Synthesis. This field study demonstrates that dark biofilms are a high‐quality resource pool for invertebrates in river ecosystems that is often overlooked. Similar to light biofilms, dark biofilms provide physiologically important FA and bacterial FA for stream invertebrates. However, these high‐quality resources are threatened by increasing human disturbances to river ecosystems. Future research is required to better understand how the quality of both dark and light biofilms changes in response to human disturbance, and how this affects dietary energy transfer to upper trophic consumers, including fish and humans. [ABSTRACT FROM AUTHOR]

The River Continuum Concept implies that consumers in headwater streams have greater dietary access to terrestrial basal resources, but recent studies have highlighted the dietary importance of high-quality algae. Algae provide consumers with physiologically important omega-3 (n-3) polyunsaturated fatty acids (PUFA), particularly eicosapentaenoic acid (EPA). However, terrestrial plants and most benthic stream algae lack the long-chain (LC) n-3 PUFA docosahexaenoic acid (DHA, 22:6n-3), which is essential for neural development in fish and other vertebrates. We sampled subalpine streams to investigate how the PUFA composition of neural (brain and eyes), muscle, and liver tissues of freshwater fish is related to their potential diets (macroinvertebrates, epilithon, fresh and conditioned terrestrial leaves). The PUFA composition of consumers was more similar to epilithon than to terrestrial leaves. Storage lipids of eyes most closely resembled dietary PUFA (aquatic invertebrates and algae). However, DHA and arachidonic acid (ARA, 20:4n-6) were not directly available in the diet but abundant in organs. This implies that algal PUFA were selectively retained or were produced internally via enzymatic PUFA conversion by aquatic consumers. This field study demonstrates the nutritional importance of algal PUFA for neural organs in aquatic consumers of headwater regions. [ABSTRACT FROM AUTHOR]

The article reports that fatty acids (FA) are a major source of nutrients and energy in aquatic food webs, as well as serving as the main components of all cell membranes. Topics include increasing anthropogenic impacts are predicted to selectively alter the production of these critical compounds, and the comprehensive assessment of these potential effects has synthesized and systematically explored differences in the abundance and distribution of FA in fish.

According to the River Continuum Concept, headwater streams are richer in allochthonous (e.g. terrestrial leaves) than autochthonous (e.g. algae) sources of organic matter for consumers. However, compared to algae, leaf litter is of lower food quality, particularly ω‐3 polyunsaturated fatty acids (n‐3 PUFA), and would constrain the somatic growth, maintenance, and reproduction of stream invertebrates. It may be thus assumed that shredders, such as Gammarus, receive lower quality diets than grazers, e.g. Ecdyonurus, that typically feed on algae.The objective of this study was to assess the provision of dietary PUFA from leaf litter and algae to the shredder Gammarus and the grazer Ecdyonurus. Three different diets (algae, terrestrial leaves, and an algae–leaf litter mix) were supplied to these macroinvertebrates in a flume experiment for 2 weeks. To differentiate how diet sources were retained in these consumers, algae were isotopically labelled with 13C.Both consumers became enriched with 13C in all treatments, demonstrating that both assimilated algae. For Gammarus, n‐3 PUFA increased, whereas n‐6 PUFA stayed constant. By contrast, the n‐3 PUFA content of Ecdyonurus decreased as a consequence of declining algal supply.Results from compound‐specific stable isotope analysis provided evidence that the long‐chain n‐3 PUFA eicosapentaenoic acid (EPA) in both consumers was more enriched in 13C than the short‐chain n‐3 PUFA α‐linolenic acid, suggesting that EPA was taken up directly from algae and not from heterotrophic biofilms on leaf litter. Both consumers depended on algae as their carbon and EPA source and retained their EPA from high‐quality algae. [ABSTRACT FROM AUTHOR]

• Fatty acid was used to trace carbon sources and evaluate food quality in streams. • Nutrient addition increased autochthonous carbon and food quality in food webs. • Trophic linkages between biofilms and grazers were determined by biofilm EPA. • Nutrient addition combined with riparian shading strengthened the trophic linkages. Eutrophication induced by excessive inputs of nutrient is one of the main stressors in aquatic ecosystems. Deforestation in riparian zones alter riparian shading, which together with eutrophication is expected to exert a complex control over stream food webs. We manipulated two levels of riparian shading (open canopy vs. shading canopy) and nutrient supply (ambient vs. nutrient addition) in three headwater streams to investigate the individual and combined effects of eutrophication and loss of riparian shading on carbon sources and nutritional quality of biofilms, and the subsequent trophic effects on macroinvertebrate grazers. Nutrient enrichment increased the autochthonous carbon (i.e., algae especially diatoms) indicated by fatty acid (FA) biomarkers within biofilms and grazers. The nutritional quality indicated by eicosapentaenoic acid (EPA) content of biofilms was increased with nutrient enrichment and more so with the combined effect of an increase in riparian shading, consequently leading to an increase in the nutritional quality, density, and biomass of grazers. In particular, the trophic linkages between biofilms and grazers were mainly influenced by EPA concentration in the biofilms, and strengthened with the combined effects of riparian shading and additional nutrients. Our study emphasizes the nutritional significance of EPA for consumers at higher trophic levels and proposes its potential as an indicator for monitoring the health of aquatic ecosystems. [Display omitted] [ABSTRACT FROM AUTHOR]

Streams and rivers provide crucial sources of long-chain polyunsaturated fatty acids ( LC- PUFA, a subset of PUFA with ≥20 C in their acyl chains) for freshwater and terrestrial animals. LC- PUFA are primarily produced by particular algal taxa and are subsequently transferred to fish and higher consumers, including humans. Although research on the trophic transfer of LC- PUFA in streams is very limited, studies on the PUFA composition of organisms at various trophic levels within stream food webs are well documented. Here, we systematically analyze the literature to synthesise LC- PUFA distribution and retention in stream ecosystems., In general, stream food webs are highly enriched in omega-3 LC- PUFA, in particular eicosapentaenoic acid ( EPA, 20:5n-3) and docosahexaenoic acid ( DHA, 22:6n-3). Algae from temperate rivers had more EPA compared with algae from subtropical rivers. Stream invertebrates from both temperate and subtropical rivers preferentially retain algal EPA and their PUFA tend to vary with algal PUFA. Invertebrates may be capable of regulating their fatty acid ( FA) composition, but this ability is limited, to the extent that they must obtain physiologically important LC- PUFA primarily from algae., DHA is selectively and highly retained in all fish taxa irrespective of their feeding sites (i.e. temperate or subtropical rivers). However, DHA content is variable among stream fish taxa and is likely linked to resource utilisation and their endogenous metabolism. Freshwater salmonids have a very high DHA content, which suggests they preferentially retain this FA or biosynthesise it from EPA. The PUFA profiles of other fish taxa more closely reflect their dietary PUFA supply. However, so far very few studies have considered the differences in the PUFA biosynthesis of fish taxa in rivers or compared the dietary PUFA effect on different fish taxa., The supply of LC- PUFA from streams and rivers is known to be affected by altered light availability, greater nutrient inputs and other anthropogenic activities, and any decrease in algal LC- PUFA production will negatively affect the consumers that are nutritionally dependent on these molecules. Further investigations are needed to understand how environmental changes affect LC- PUFA production and retention in stream ecosystems, and how to protect streams from threats to this essential ecosystem service. [ABSTRACT FROM AUTHOR]

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Few studies measure multiple ecological tracers in individual organisms, thus limiting our ability to differentiate among organic matter source pathways and understand consequences of dietary variation and the use of external subsidies in complex food webs. We combined two tracers, stable isotope (SI) ratios and fatty acids (FA), to investigate linkages among ecological compartments (water column, benthos, riparian zone) in food webs in waterholes of a dryland river network, the Border Rivers in southwestern Queensland, Australia. Comprehensive analyses of sources (plankton, periphyton, leaf litter, riparian grasses) and animals (benthic insects, mollusks, large crustaceans, fishes) for SI and FA showed that all three zones contribute to animal biomass, depending on species and life stage. Large fishes derived a subsidy from the riparian/floodplain zone, likely through the consumption of terrestrial and semi-aquatic insects and prawns that fed on detritivorous insects. Importantly, post-larval bony bream (Nematalosa erebi) and golden perch (Macquaria ambigua) were tightly connected to the water column, as evidenced by 13C-depleted, 15Nenriched isotope ratios and a high content of plankton-derived polyunsaturated fatty acids (eicosapentaenoic acid [EPA; 20:5ω3] and docosahexaenoic acid [DHA; 22:6ω3]). These observations were consistent with expectations from nutritional requirements of fish early life stages and habitat changes associated with maturity. These results highlight the importance of high-quality foods during early development of fishes, and show that attempting to attribute food-web production to a single source pathway overlooks important but often subtle subsidies that maintain viable populations. A complete understanding of food-web dynamics must consider both quantity and quality of different available organic matter sources. This understanding can be achieved with a combined SI and FA approach, but more controlled dietary studies are needed to estimate how FA profiles are modified by animals when consuming a diverse range of diets of variable quality. [ABSTRACT FROM AUTHOR]

Gammarid amphipods are a crucial link connecting primary producers with secondary consumers, but little is known about their nutritional ecology. Here we asked how starvation and subsequent feeding on different nutritional quality algae influences fatty acid retention, compound-specific isotopic carbon fractionation, and biosynthesis of ω-3 and ω-6 polyunsaturated fatty acids (PUFA) in the relict gammarid amphipod Pallaseopsis quadrispinosa. The fatty acid profiles of P. quadrispinosa closely matched with those of the dietary green algae after only seven days of refeeding, whereas fatty acid patterns of P. quadrispinosa were less consistent with those of the diatom diet. This was mainly due to P. quadrispinosa suffering energy limitation in the diatom treatment which initiated the metabolization of 16:1ω7 and partly 18:1ω9 for energy, but retained high levels of eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA) similar to those found in wild-caught organisms. Moreover, α-linolenic acid (ALA) from green algae was mainly stored and not allocated to membranes at high levels nor biosynthesized to EPA. The arachidonic acid (ARA) content in membrane was much lower than EPA and P. quadrispinosa was able to biosynthesize long-chain ω-6 PUFA from linoleic acid (LA). Our experiment revealed that diet quality has a great impact on fatty acid biosynthesis, retention and turnover in this consumer. [ABSTRACT FROM AUTHOR]