Terrestrial organic matter inputs have long been thought to play an important role in aquatic food web dynamics. Results from recent whole lake 13 C addition experiments suggest terrestrial particulate organic carbon (t-POC) inputs account for a disproportionate portion of zooplankton production. For example, several studies concluded that although t-POC only represented Ϸ20% of the flux of particulate carbon available to herbivorous zooplankton, this food source accounted for Ϸ50% of the C incorporated by zooplankton. We tested the direct dietary impact of t-POC (from the leaves of riparian vegetation) and various phytoplankton on Daphnia magna somatic growth, reproduction, growth efficiency, and lipid composition. By itself, t-POC was a very poor quality resource compared to cryptophytes, diatoms, and chlorophytes, but t-POC had similar food quality compared to cyanobacteria. Small additions of high quality Cryptomonas ozolinii to t-POC-dominated diets greatly increased Daphnia growth and reproduction. When offered alone, t-POC resulted in a Daphnia growth efficiency of 5 ؎ 1%, whereas 100% Cryptomonas and Scenedesmus obliquus diets resulted in growth efficiencies of 46 ؎ 8% (؎ SD) and 36 ؎ 3%, respectively. When offered in a 50:50 mixed diet with Cryptomonas or Scenedesmus, the t-POC fraction resulted in a partial growth efficiency of 22 ؎ 9% and 15 ؎ 6%, respectively. Daphnia that obtained 80% of their available food from t-POC assimilated 84% of their fatty acids from the phytoplankton component of their diet. Overall, our results suggest Daphnia selectively allocate phytoplankton-derived POC and lipids to enhance somatic growth and reproduction, while t-POC makes a minor contribution to zooplankton production.Daphnia ͉ fatty acids ͉ nutritional ecology ͉ planktonic food web

Long-chain polyunsaturated fatty acids (LC-PUFA) are critical for the health of aquatic and terrestrial organisms; therefore, understanding the production, distribution, and abundance of these compounds is very important. Although the dynamics of LC-PUFA production and distribution in aquatic environments has been well documented, a systematic and comprehensive comparison to LC-PUFA in terrestrial environments has not been rigorously investigated. Here we use a data synthesis approach to compare and contrast fatty acid profiles of 369 aquatic and terrestrial organisms. Habitat and trophic level were interacting factors that determined the proportion of individual omega-3 (n-3) or omega-6 (n-6) PUFA in aquatic and terrestrial organisms. Higher total n-3 content compared with n-6 PUFA and a strong prevalence of the n-3 PUFA eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA) characterized aquatic versus terrestrial organisms. Conversely, terrestrial organisms had higher linoleic acid (LNA) and alpha-linolenic acid (ALA) contents than aquatic organisms; however, the ratio of ALA:LNA was higher in aquatic organisms. The EPA + DHA content was higher in aquatic animals than terrestrial organisms, and increased from algae to invertebrates to vertebrates in the aquatic environment. An analysis of covariance revealed that fatty acid composition was highly dependent on the interaction between habitat and trophic level. We conclude that freshwater ecosystems provide an essential service through the production of n-3 LC-PUFA that are required to maintain the health of terrestrial organisms including humans.

Abstract1. Many freshwater systems receive substantial inputs of terrestrial organic matter.Terrestrially derived dissolved organic carbon (t-DOC) inputs can modify light availability, the spatial distribution of primary production, heat, and oxygen in aquatic systems, as well as inorganic nutrient bioavailability. It is also well-established that some terrestrial inputs (such as invertebrates and fruits) provide highquality food resources for consumers in some systems.2. In small to moderate-sized streams, leaf litter inputs average approximately three times greater than the autochthonous production. Conversely, in oligo/mesotrophic lakes algal production is typically five times greater than the available flux of allochthonous basal resources.3. Terrestrial particulate organic carbon (t-POC) inputs to lakes and rivers are comprised of 80%-90% biochemically recalcitrant lignocellulose, which is highly resistant to enzymatic breakdown by animal consumers. Further, t-POC and heterotrophic bacteria lack essential biochemical compounds that are critical for rapid growth and reproduction in aquatic invertebrates and fishes. Several studies have directly shown that these resources have very low food quality for herbivorous zooplankton and benthic invertebrates. 4.Much of the nitrogen assimilated by stream consumers is probably of algal origin, even in systems where there appears to be a significant terrestrial carbon contribution. Amino acid stable isotope analyses for large river food webs indicate that most upper trophic level essential amino acids are derived from algae. Similarly, profiles of essential fatty acids in consumers show a strong dependence on the algal food resources.5. Primary production to respiration ratios are not a meaningful index to assess consumer allochthony because respiration represents an oxidised carbon flux that cannot be utilised by animal consumers. Rather, the relative importance of allochthonous subsidies for upper trophic level production should be addressed by considering the rates at which terrestrial and autochthonous resources are consumed and the growth efficiency supported by this food.6. Ultimately, the biochemical composition of a particular basal resource, and not just its quantity or origin, determines how readily this material is incorporated into upper trophic level consumers. Because of its highly favourable biochemical composition and greater availability, we conclude that microalgal production supports most animal production in freshwater ecosystems. | WHY DOES ALLOCHTHONY MATTER?A better understanding of where and how allochthony modifies aquatic food web processes will improve our ability to predict how land-use and climate change affect organic carbon export from watersheds to lakes and rivers and how this matter influences upper trophic level production in aquatic systems. If invertebrate and fish consumers in rivers and lakes are strongly subsidised by allochthonous carbon inputs, then watersheds and especially riparian zone management will potentially have as mu...

SUMMARY1. While many streams and rivers are dominated by terrestrial inputs of organic carbon, algae are an important trophic base for stream food webs. However, the nutritional importance of algae for stream invertebrates only recently has been highlighted. Algae are acknowledged as higher quality food than terrestrial organic matter for the growth and reproduction of invertebrates. In part, this is because of higher algal polyunsaturated fatty-acid (PUFA) content. Here, we review the important influence of algal food quality, as assessed by PUFA, in stream food webs. 2. Current field investigations have mainly focused on the fatty-acid dynamics of macroinvertebrates, and indicate that algal eicosapentaenoic acid (EPA), a-linolenic acid (ALA) and linoleic acids (LIN) are present in all macroinvertebrates. However, fungal and bacterial tracers have also been observed in a range of macroinvertebrates. The omega-3 (x3)/omega-6 (x6) ratio >1 in most macroinvertebrates strongly indicates that dietary energy of algae is highly retained in stream food webs. Interspecific differences in PUFA composition seem to be affected by dietary PUFA and consumer physiology. 3. Some studies have suggested that besides dietary EPA, the shorter chain C18 PUFA LIN and ALA also can improve growth and reproduction of stream invertebrate consumers. Some macroinvertebrates may preferentially retain or synthesise long-chain PUFA from C18 PUFA when experiencing low-quality food. However, this process is controversial since other species have shown very limited ability to synthesise long-chain PUFA. 4. Algal PUFA composition is strongly influenced by abiotic factors, particularly light, nutrients, and temperature. Human disturbance (i.e. riparian vegetation removal and nutrient inputs) on algal PUFA content and their consequent effects on macroinvertebrates and fish clearly warrant further scientific attention. Controlled feeding trials and manipulative studies are required to measure PUFA conversion capacities and reproductive investment of stream macroinvertebrates under different food quality conditions, which will provide insights into how freshwater species can cope with different nutritional food conditions due to human disturbance and climate change.

Zooplankton transfer ecologically important fatty acids (FA) from their diets to upper trophic levels. We used diet-switching experiments with 13 C-labeled food sources to determine the time scale at which dietary uptake is manifested in the FA profi les of Daphnia magna . Daphnia dramatically shifted their FA composition in response to diet change within only four days, however Daphnia switched from a high quality (i.e. Cryptomonas ) to a moderate quality ( Scenedesmus ) diet retained the most physiologically important FA from their original diet source even after 14 days. In particular, Daphnia exhibited long-term retention of eicosapentaenoic (EPA; 20:5 ω 3) and arachidonic acid (ARA; 20:4 ω 6) when switched from Cryptomonas to Scenedesmus . Similarly, when switched from Scenedesmus to Cryptomonas , Daphnia took up a high proportion of EPA and ARA after only two days. Th e phospholipid fatty acid (PLFA) fraction in Daphnia was preferentially enriched with stearic (18:0), oleic (18:1 ω 9), and linoleic acid (LIN; 18:2 ω 6). In contrast with studies of marine copepods, dietary FA also strongly aff ected the PLFA composition (structural lipids) of Daphnia . Results of δ 13 C signatures of individual FA provided evidence of elongation and desaturation of α -linolenic (ALA; 18:3 ω 3) or stearidonic acid (SDA; 18:4 ω 3) to EPA 10 days after a diet switch to EPA-defi cient Scenedesmus . Diff erences in the ARA content of Daphnia fed Cryptomonas and Scenedesmus suggest Daphnia consuming Cryptomonas synthesized ARA via retroconversion of ω 6-docosapentaenoic acid ( ω 6-DPA; 22:5 ω 6). Daphnia preferentially accumulate and retain, as well as bioconvert, those FA that are also most physiologically important for fi sh production. Our results also indicate Daphnia FA composition responds to their diet on a short temporal scale and analyses of lipid biomarkers in zooplankton provide strong insights into the food sources that support their production.

Polyunsaturated fatty acids (PUFA), especially long-chain (i.e., ≥20 carbons) polyunsaturated fatty acids (LC-PUFA), are fundamental to the health and survival of marine and terrestrial organisms. Therefore, it is imperative that we gain a better understanding of their origin, abundance, and transfer between and within these ecosystems. We evaluated the natural variation in PUFA distribution and abundance that exists between and within these ecosystems by amassing and analyzing, using multivariate and analysis of variance (ANOVA) methods, >3000 fatty acid (FA) profiles from marine and terrestrial organisms. There was a clear dichotomy in LC-PUFA abundance between organisms in marine and terrestrial ecosystems, mainly driven by the C18 PUFA in terrestrial organisms and omega-3 (n-3) LC-PUFA in marine organisms. The PUFA content of an organism depended on both its biome (marine vs terrestrial) and taxonomic group. Within the marine biome, the PUFA content varied among taxonomic groups. PUFA content of marine organisms was dependent on both geographic zone (i.e., latitude, and thus broadly related to temperature) and trophic level (a function of diet). The contents of n-3 LC-PUFA were higher in polar and temperate marine organisms than those from the tropics. Therefore, we conclude that, on a per capita basis, high latitude marine organisms provide a disproportionately large global share of these essential nutrients to consumers, including terrestrial predators. Our analysis also hints at how climate change, and other anthropogenic stressors, might act to negatively impact the global distribution and abundance of n-3 LC-PUFA within marine ecosystems and on the terrestrial consumers that depend on these subsidies.

Mountain lakes are often situated in protected natural areas, a feature that leads to their role as sentinels of global environmental change. Despite variations in latitude, mountain lakes share many features, including their location in catchments with steep topographic gradients, cold temperatures, high incident solar and ultraviolet radiation (UVR), and prolonged ice and snow cover. These characteristics, in turn, affect mountain lake ecosystem structure, diversity, and productivity. The lakes themselves are mostly small and shallow, and up until recently, have been characterized as oligotrophic. This paper provides a review and update of the growing body of research that shows that sediments in remote mountain lakes archive regional and global environmental changes, including those linked to climate change, altered biogeochemical cycles, and changes in dust composition and deposition, atmospheric fertilization, and biological manipulations. These archives provide an important record of global environmental change that pre-dates typical monitoring windows. Paleolimnological research at strategically selected lakes has increased our knowledge of interactions among multiple stressors and their synergistic effects on lake systems. Lakes from transects across steep climate (i.e., temperature and effective moisture) gradients in mountain regions show how environmental change alters lakes in close proximity, but at differing climate starting points. Such research in particular highlights the impacts of melting glaciers on mountain lakes. The addition of new proxies, including DNAbased techniques and novel stable isotopic analyses, provides a gateway to addressing novel research questions about global environmental change. Recent advances in remote sensing and continuous, high-frequency, limnological measurements will improve spatial and temporal resolution and help to add records to spatial gaps including tropical and southern latitudes.

We explored how dietary bacteria affect the life history traits and biochemical composition of Daphnia magna, using three bacteria taxa with very different lipid composition. Our objectives were to (1) examine whether and how bacteria-dominated diets affect Daphnia survival, growth, and fecundity, (2) see whether bacteria-specific fatty acid (FA) biomarkers accrued in Daphnia lipids, and (3) explore the quantitative relationship between bacteria availability in Daphnia diets and the amounts of bacterial FA in their lipids. Daphnia were fed monospecific and mixed diets of heterotrophic (Micrococcus luteus) or methanotrophic bacteria (Methylomonas methanica and Methylosinus trichosporium) and two phytoplankton species (Cryptomonas ozolinii and Scenedesmus obliquus). Daphnia neonates fed pure bacteria diets died after 6-12 days and produced no viable offspring, whereas those fed pure phytoplankton diets had high survival, growth, and reproduction success. Daphnia fed a mixed diet with 80% M. luteus and 20% of either phytoplankton had high somatic growth, but low reproduction. Conversely, Daphnia fed mixed diets including 80% of either methane-oxidizing bacteria and 20% Cryptomonas had high reproduction rates, but low somatic growth. All Daphnia fed mixed bacteria and phytoplankton diets had strong evidence of both bacteria-and phytoplankton-specific FA biomarkers in their lipids. FA mixing model calculations indicated that Daphnia that received 80% of their carbon from bacteria assimilated 46 ± 25% of their FA from this source. A bacteria-phytoplankton gradient experiment showed a strong positive correlation between the proportions of the bacterial FA in the Daphnia and their diet, indicating that bacterial utilization can be traced in this keystone consumer using FA biomarkers.