Quantifying the energetic cost of food quality constraints on resting metabolism to integrate nutritional and metabolic ecology.

Consumer metabolism controls the energy uptake from the environment and its allocation to biomass production. In natural ecosystems, available energy in food often fails to predict biomass production which is also (co)limited by the relative availability of various dietary compounds. To date, the link between energy metabolism and the effects of food chemical composition on biomass production remains elusive. Here, we measured the resting metabolic rate (RMR) of Daphnia magna along ontogeny when undergoing various (non-energetic) nutritional constraints. All types of dietary (co)limitations (Fatty acids, Sterols, Phosphorus) induced an increase in mass-specific RMR up to 128% between highest and lowest quality diets. We highlight a strong negative correlation between RMR and growth rate indicating RMR as a promising predictor of consumer growth rate. We argue that quantifying the energetic cost imposed by food quality on individual RMR may constitute a common currency enabling the integration of nutritional and metabolic ecology.

U-shaped response Unifies views on temperature dependency of stoichiometric requirements.

Temperature and nutrient availability, which are major drivers of consumer performance, are dramatically affected by global change. To date, there is no consensus on whether warming increases or decreases consumer needs for dietary carbon (C) relatively to phosphorus (P), thus hindering predictions of secondary production responses to global change. Here, we investigate how the dietary C:P ratio optimising consumer growth (TERC:P : Threshold Elemental Ratio) changes along temperature gradients by combining a temperature-dependent TERC:P model with growth experiments on Daphnia magna. Both lines of evidence show that the TERC:P response to temperature is U-shaped. This shape indicates that consumer nutrient requirements can both increase or decrease with increasing temperature, thus reconciling previous contradictive observations into a common framework. This unified framework improves our capacity to forecast the combined effects of nutrient cycle and climatic alterations on invertebrate production.

Simulating rewetting events in intermittent rivers and ephemeral streams: A global analysis of leached nutrients and organic matter.

Climate change and human pressures are changing the global distribution and the extent of intermittent rivers and ephemeral streams (IRES), which comprise half of the global river network area. IRES are characterized by periods of flow cessation, during which channel substrates accumulate and undergo physico-chemical changes (preconditioning), and periods of flow resumption, when these substrates are rewetted and release pulses of dissolved nutrients and organic matter (OM). However, there are no estimates of the amounts and quality of leached substances, nor is there information on the underlying environmental constraints operating at the global scale. We experimentally simulated, under standard laboratory conditions, rewetting of leaves, riverbed sediments, and epilithic biofilms collected during the dry phase across 205 IRES from five major climate zones. We determined the amounts and qualitative characteristics of the leached nutrients and OM, and estimated their areal fluxes from riverbeds. In addition, we evaluated the variance in leachate characteristics in relation to selected environmental variables and substrate characteristics. We found that sediments, due to their large quantities within riverbeds, contribute most to the overall flux of dissolved substances during rewetting events (56%-98%), and that flux rates distinctly differ among climate zones. Dissolved organic carbon, phenolics, and nitrate contributed most to the areal fluxes. The largest amounts of leached substances were found in the continental climate zone, coinciding with the lowest potential bioavailability of the leached OM. The opposite pattern was found in the arid zone. Environmental variables expected to be modified under climate change (i.e. potential evapotranspiration, aridity, dry period duration, land use) were correlated with the amount of leached substances, with the strongest relationship found for sediments. These results show that the role of IRES should be accounted for in global biogeochemical cycles, especially because prevalence of IRES will increase due to increasing severity of drying events.

A microcalorimetric approach for investigating stoichiometric constraints on the standard metabolic rate of a small invertebrate.

Understanding the determinants of metabolism is a core ecological topic since it permits to link individual energetic requirements to the ecology of communities and ecosystems. Yet, besides temperature, metabolic responses to environmental factors remain poorly understood. For example, it is commonly assumed that dietary stoichiometric constraints increase metabolism of small invertebrates despite scarce experimental support. Here, we used microcalorimetric measurements to determine the standard metabolic rate (SMR) of Daphnia magna fed stoichiometrically balanced (C/P: 166) or imbalanced (C/P: 1439). Daphnids fed imbalanced maintained their stoichiometric homeostasis within narrow boundaries. However, they consistently increased their SMR while decreasing their growth rate. Our measurements demonstrate that homeostatic regulation implies higher metabolic costs, thereby reducing available energy for growth. We demonstrate that microcalorimetry is a powerful and precise tool for measuring small-sized organisms' metabolic rate, thus opening promising perspectives for understanding how environmental factors, such as nutritional constraints, affect organismal metabolism.

Litter identity mediates predator impacts on the functioning of an aquatic detritus-based food web.

During past decades, several mechanisms such as resource quality and habitat complexity have been proposed to explain variations in the strength of trophic cascades across ecosystems. In detritus-based headwater streams, litter accumulations constitute both a habitat and a resource for detritivorous macroinvertebrates. Because litter edibility (which promotes trophic cascades) is usually inversely correlated with its structural complexity (which weakens trophic cascades), there is a great scope for stronger trophic cascades in litter accumulations that are dominated by easily degradable litter species. However, it remains unclear how mixing contrasting litter species (conferring both habitat complexity and high quality resource) may influence top-down controls on communities and processes. In enclosures exposed in a second-order stream, we manipulated litter species composition by using two contrasting litter (alder and oak), and the presence-absence of a macroinvertebrate predator (Cordulegaster boltonii larvae), enabling it to effectively exert predation pressure, or not, on detritivores (consumptive versus non-consumptive predation effects). Leaf mass loss, detritivore biomass and community structure were mostly controlled independently by litter identity and mixing and by predator consumption. However, the strength of predator control was mediated by litter quality (stronger on alder), and to a lesser extent by litter mixing (weaker on mixed litter). Refractory litter such as oak leaves may contribute to the structural complexity of the habitat for stream macroinvertebrates, allowing the maintenance of detritivore communities even when strong predation pressure occurs. We suggest that considering the interaction between top-down and bottom-up factors is important when investigating their influence on natural communities and ecosystem processes in detritus-based ecosystems.

Photodegradation of disposable polypropylene face masks: Physicochemical properties of debris and implications for the toxicity of mask-carried river biofilms.

COVID-19 outbreak led to a massive dissemination of protective polypropylene (PP) face masks in the environment, posing a new environmental risk amplified by mask photodegradation and fragmentation. Masks are made up of a several kilometres long-network of fibres with diameter from a few microns to around 20 µm. After photodegradation, these fibres disintegrate, producing water dispersible debris. Electrokinetics and particle stability observations support that photodegradation increases/decreases the charge/hydrophobicity of released colloidal fragments. This change in hydrophobicity is related to the production of UV-induced carbonyl and hydroxyl reactive groups detectable after a few days of exposure. Helical content, surface roughness and specific surface area of mask fibres are not significantly impacted by photodegradation. Fragmentation of fibres makes apparent, at the newly formed surfaces, otherwise-buried additives like TiO2 nanoparticles and various organic components. Mortality of gammarids is found to increase significantly over time when fed with 3 days-UV aged masks that carry biofilms grown in river, which is due to a decreased abundance of microphytes therein. In contrast, bacteria abundance and microbial community composition remain unchanged regardless of mask degradation. Overall, this work reports physicochemical properties of pristine and photodegraded masks, and ecosystemic functions and ecotoxicity of freshwater biofilms they can carry.

Benthic algae stimulate leaf litter decomposition in detritus-based headwater streams: a case of aquatic priming effect?

In detritus-based ecosystems, autochthonous primary production contributes very little to the detritus pool. Yet primary producers may still influence the functioning of these ecosystems through complex interactions with decomposers and detritivores. Recent studies have suggested that, in aquatic systems, small amounts of labile carbon (C) (e.g., producer exudates), could increase the mineralization of more recalcitrant organic-matter pools (e.g., leaf litter). This process, called priming effect, should be exacerbated under low-nutrient conditions and may alter the nature of interactions among microbial groups, from competition under low-nutrient conditions to indirect mutualism under high-nutrient conditions. Theoretical models further predict that primary producers may be competitively excluded when allochthonous C sources enter an ecosystem. In this study, the effects of a benthic diatom on aquatic hyphomycetes, bacteria, and leaf litter decomposition were investigated under two nutrient levels in a factorial microcosm experiment simulating detritus-based, headwater stream ecosystems. Contrary to theoretical expectations, diatoms and decomposers were able to coexist under both nutrient conditions. Under low-nutrient conditions, diatoms increased leaf litter decomposition rate by 20% compared to treatments where they were absent. No effect was observed under high-nutrient conditions. The increase in leaf litter mineralization rate induced a positive feedback on diatom densities. We attribute these results to the priming effect of labile C exudates from primary producers. The presence of diatoms in combination with fungal decomposers also promoted decomposer diversity and, under low-nutrient conditions, led to a significant decrease in leaf litter C:P ratio that could improve secondary production. Results from our microcosm experiment suggest new mechanisms by which primary producers may influence organic matter dynamics even in ecosystems where autochthonous primary production is low.

Impaired leaf litter processing in acidified streams : learning from microbial enzyme activities.

Anthropogenic acidification in headwater streams is known to affect microbial assemblages involved in leaf litter breakdown. Far less is known about its potential effects on microbial enzyme activities. To assess the effects of acidification on microbial activities associated with decaying leaves, a 70-day litter bag experiment was conducted in headwater streams at six sites across an acidification gradient. The results revealed that microbial leaf decomposition was strongly and negatively correlated with total Al concentrations (r = -0.99, p < 0.001) and positively correlated with Ca(2+) concentrations (r = 0.94, p = 0.005) and pH (r = 0.93, p = 0.008). Denaturing gradient gel electrophoresis analyses showed that microbial assemblages differed between non-impacted and impacted sites, whereas fungal biomass associated with decaying leaves was unaffected. The nutrient content of leaf detritus and ecoenzymatic activities of carbon (C), nitrogen (N) and phosphorus (P) acquisition revealed that N acquisition was unaltered, while P acquisition was significantly reduced across the acidification gradient. The P content of leaf litter was negatively correlated with total Al concentrations (r = -0.94, p < 0.01) and positively correlated with decomposition rates (r = 0.95, p < 0.01). This potential P limitation of microbial decomposers in impacted sites was confirmed by the particularly high turnover activity for phosphatase and imbalanced ratios between the ecoenzymatic activities of C and P acquisition. The toxic form of Al has well-known direct effects on aquatic biota under acidic conditions, but in this study, Al was found to also potentially affect microbially mediated leaf processing by interfering with the P cycle. These effects may in turn have repercussions on higher trophic levels and whole ecosystem functioning.

A 34-year survey under phosphorus decline and warming: Consequences on stoichiometry and functional trait composition of freshwater macroinvertebrate communities.

Worldwide, freshwater systems are subjected to increasing temperatures and nutrient changes. Under phosphorus and nitrogen enrichment consumer communities are often thought to shift towards fast-growing and P-rich taxa, supporting the well-known link between growth rate and body stoichiometry. While these traits are also favoured under warming, the temperature effect on stoichiometry is less clear. As recently shown, there is a general link between functional traits and body stoichiometry, which makes the integration of stoichiometric traits a promising tool to help understanding the mechanisms behind taxonomic and functional community responses to nutrient changes and/or warming. Yet, such approaches have been scarcely developed at community level and on a long-term perspective. In this study, we investigated long-term responses in stoichiometry and functional trait composition of macroinvertebrate communities to nutrient changes (decreasing water P; increasing water N:P) and warming over a 34-year period in the Middle Loire River (France), testing the potentially opposing responses to these drivers. Both drivers should cause shifts in species composition, which will alter the overall community stoichiometry and functional composition following assumptions from ecological stoichiometry theory. We found that the macroinvertebrate community shifted towards P-poor taxa, causing significant trends in overall community stoichiometry which indicates long-term changes in the nutrient pool provided by these consumers (i.e. decrease in %N and %P, increase in N:P). Further, while the former high-P conditions favoured traits associated to detritus feeding and fast development (i.e. small maximum body size, short life duration), recent conditions favoured predators and slow-developing taxa. These results suggest nutrients to be a more important driver than temperature over this period. By providing a pivotal link between environmental changes and functional trait composition of communities, approaches based on stoichiometric traits offer sound perspectives to investigate ecological relationships between multiple drivers operating at various scales and ecosystem functioning.

Nitrogen to phosphorus ratio shapes the bacterial communities involved in cellulose decomposition and copper contamination alters their stoichiometric demands.

All living organisms theoretically have an optimal stoichiometric nitrogen: phosphorus (N: P) ratio, below and beyond which their growth is affected, but data remain scarce for microbial decomposers. Here, we evaluated optimal N: P ratios of microbial communities involved in cellulose decomposition and assessed their stability when exposed to copper Cu(II). We hypothesized that (1) cellulose decomposition is maximized for an optimal N: P ratio; (2) copper exposure reduces cellulose decomposition and (3) increases microbial optimal N: P ratio; and (4) N: P ratio and copper modify the structure of microbial decomposer communities. We measured cellulose disc decomposition by a natural inoculum in microcosms exposed to a gradient of N: P ratios at three copper concentrations (0, 1 and 15 µM). Bacteria were most probably the main decomposers. Without copper, cellulose decomposition was maximized at an N: P molar ratio of 4.7. Contrary to expectations, at high copper concentration, the optimal N: P ratio (2.8) and the range of N: P ratios allowing decomposition were significantly reduced and accompanied by a reduction of bacterial diversity. Copper contamination led to the development of tolerant taxa probably less efficient in decomposing cellulose. Our results shed new light on the understanding of multiple stressor effects on microbial decomposition in an increasingly stoichiometrically imbalanced world.

Lessons from linking bio- and ecological traits to stoichiometric traits in stream macroinvertebrates.

Ecologists rely on various functional traits when investigating the functioning of ecological systems and its responses to global changes. Changing nutrient levels, for example, can affect taxa expressing different trait combinations in various ways, e.g., favoring small, fast-growing species under high phosphorus conditions. Stoichiometric traits, describing the elemental composition of organism body tissues, can help in understanding the mechanisms behind such functional shifts. So far, mainly life-history traits have been related to body stoichiometry (e.g., the growth rate hypothesis) on a limited number of taxa, and there is little knowledge of the general link between stoichiometric and other functional traits on a taxonomically large scale. Here, we highlight this link in the freshwater macroinvertebrates, testing predictions from underlying trait-based and Ecological Stoichiometry Theory (EST) in >200 taxa belonging to eight larger taxonomic groups. We applied a series of multivariate analyses on six of their stoichiometric traits (%C, %N, %P, C:N, C:P, and N:P) and 23 biological and ecological traits. We found significant relationships between stoichiometric traits and other types of traits when analyzing single-trait and multi-trait profiles. Patterns found within traits related to organism development or nutrient cycling were in line with our assumptions based on EST, e.g., traits describing predators were associated with high %N; traits suggesting a fast development (small maximum body size and high molting frequency) with high %P. Associations between ecological traits and body stoichiometry could be explained by the longitudinal stream gradient: Taxa preferring headwater habitats (i.e., high altitude, coarse substrate, and cold temperature) exhibited high %N and %P. Demonstrating the link between stoichiometric and both bio- and ecological traits on a large diversity of taxa underlines the potential of integrating stoichiometric traits into ecological analyses to improve our understanding of taxonomic and functional responses of communities-and ecosystems-to changing environmental conditions worldwide.

Organic-matter decomposition as a bioassessment tool of stream functioning: A comparison of eight decomposition-based indicators exposed to different environmental changes.

Organic-matter decomposition has long been proposed as a tool to assess stream functional integrity, but this indicator largely depends on organic-matter selection. We assessed eight decomposition-based indicators along two well-known environmental gradients, a nutrient-enrichment gradient (0.2-1.4 mg DIN/L) in central Portugal and an acidification gradient (pH: 4.69-7.33) in north-eastern France to identify the most effective organic-matter indicator for assessing stream functional integrity. Functional indicators included natural leaf litter (alder and oak) in 10-mm and 0.5-mm mesh bags, commercial tea (Lipton green and rooibos teas in 0.25-mm mesh bags), wood sticks (wood tongue depressors) and cotton strips. Biotic indices based on benthic macroinvertebrates (IPtIN for Portugal and IBGN for France) were calculated to compare the effectiveness of structural and functional indicators in detecting stream impairment and to assess the relationship between both types of indicators. The effectiveness of organic-matter decomposition rates as a functional indicator depended on the stressor considered and the substrate used. Decomposition rates generally identified nutrient enrichment and acidification in the most acidic streams. Decomposition rates of alder and oak leaves in coarse-mesh bags, green and rooibos teas and wood sticks were positively related with pH. Only decomposition rates of rooibos tea and wood sticks were related with DIN concentration; decomposition rates along the nutrient-enrichment gradient were confounded by differences in shredder abundance and temperature among streams. Stream structural integrity was good to excellent across streams; the IPtIN index was unrelated to DIN concentration, while the IBGN index was positively related with pH. The relationships between decomposition rates and biotic indices were loose in most cases, and only decomposition rates of alder leaves in coarse-mesh bags and green tea were positively related with the IBGN. Commercial substrates may be a good alternative to leaf litter to assess stream functional integrity, especially in the case of nutrient enrichment.

Priming effect: bridging the gap between terrestrial and aquatic ecology.

Understanding how ecosystems store or release carbon is one of ecology's greatest challenges in the 21st century. Organic matter covers a large range of chemical structures and qualities, and it is classically represented by pools of different recalcitrance to degradation. The interaction effects of these pools on carbon cycling are still poorly understood and are most often ignored in global-change models. Soil scientists have shown that inputs of labile organic matter frequently tend to increase, and often double, the mineralization of the more recalcitrant organic matter. The recent revival of interest for this phenomenon, named the priming effect, did not cross the frontiers of the disciplines. In particular, the priming effect phenomenon has been almost totally ignored by the scientific communities studying marine and continental aquatic ecosystems. Here we gather several arguments, experimental results, and field observations that strongly support the hypothesis that the priming effect is a general phenomenon that occurs in various terrestrial, freshwater, and marine ecosystems. For example, the increase in recalcitrant organic matter mineralization rate in the presence of labile organic matter ranged from 10% to 500% in six studies on organic matter degradation in aquatid ecosystems. Consequently, the recalcitrant organic matter mineralization rate may largely depend on labile organic matter availability, influencing the CO2 emissions of both aquatic and terrestrial ecosystems. We suggest that (1) recalcitrant organic matter may largely contribute to the CO2 emissions of aquatic ecosystems through the priming effect, and (2) priming effect intensity may be modified by global changes, interacting with eutrophication processes and atmospheric CO2 increases. Finally, we argue that the priming effect acts substantially in the carbon and nutrient cycles in all ecosystems. We outline exciting avenues for research, which could provide new insights on the responses of ecosystems to anthropogenic perturbations and their feedbacks to climatic changes.

Continental-scale patterns of hyper-cryptic diversity within the freshwater model taxon Gammarus fossarum (Crustacea, Amphipoda).

Traditional morphological diagnoses of taxonomic status remain widely used while an increasing number of studies show that one morphospecies might hide cryptic diversity, i.e. lineages with unexpectedly high molecular divergence. This hidden diversity can reach even tens of lineages, i.e. hyper cryptic diversity. Even well-studied model-organisms may exhibit overlooked cryptic diversity. Such is the case of the freshwater crustacean amphipod model taxon Gammarus fossarum. It is extensively used in both applied and basic types of research, including biodiversity assessments, ecotoxicology and evolutionary ecology. Based on COI barcodes of 4926 individuals from 498 sampling sites in 19 European countries, the present paper shows (1) hyper cryptic diversity, ranging from 84 to 152 Molecular Operational Taxonomic Units, (2) ancient diversification starting already 26 Mya in the Oligocene, and (3) high level of lineage syntopy. Even if hyper cryptic diversity was already documented in G. fossarum, the present study increases its extent fourfold, providing a first continental-scale insight into its geographical distribution and establishes several diversification hotspots, notably south-eastern and central Europe. The challenges of recording hyper cryptic diversity in the future are also discussed.

High food quality increases infection of Gammarus pulex (Crustacea: Amphipoda) by the acanthocephalan parasite Pomphorhynchus laevis.

Parasitism is an important process in ecosystems, but has been largely neglected in ecosystem research. However, parasites are involved in most trophic links in food webs with, in turn, a major role in community structure and ecosystem processes. Several studies have shown that higher nutrient availability in ecosystems tends to increase the prevalence of parasites. Yet, most of these studies focused on resource availability, whereas studies investigating resource quality remain scarce. In this study, we tested the impact of the quality of host food resources on infection by parasites, as well as on the consequences for the host. Three resources were used to individually feed Gammarus pulex (Crustacea: Amphipoda) experimentally infected or not infected with the acanthocephalan species Pomphorhynchus laevis: microbially conditioned leaf litter without phosphorus input (standard resource); microbially conditioned leaf litter enriched in phosphorus; and microbially conditioned leaf litter without phosphorus input but complemented with additional inputs of benthic diatoms rich in both phosphorus and eicosapentaenoic acid. During the 110 day experiment, infection rate, parasite load, host survival, and parasite-mediated behavioral traits implicated in trophic transmission were measured (refuge use, geotaxis and locomotor activity). The resources of higher quality, regardless of the infection status, reduced gammarid mortality and increased gammarid growth. In addition, higher quality resources increased the proportion of infected gammarids, and led to more cases of multi-infections. While slightly modifying the geotaxis behavior of uninfected gammarids, resource quality did not modulate the impact of parasites on host behavior. Finally, for most parameters, consumption of algal resources had a greater impact than did phosphorus-enriched leaf litter. Therefore, manipulation of resource quality significantly affected host-parasite relationships, which stressed the need for future research to investigate in natura the relationships between resource availability, resource quality and parasite prevalence.

Interactive Impacts of Silver and Phosphorus on Autotrophic Biofilm Elemental and Biochemical Quality for a Macroinvertebrate Consumer.

Autotrophic biofilms are complex and fundamental biological compartments of many aquatic ecosystems. In particular, these biofilms represent a major resource for many invertebrate consumers and the first ecological barrier against toxic metals. To date, very few studies have investigated the indirect effects of stressors on upper trophic levels through alterations of the quality of biofilms for their consumers. In a laboratory study, we investigated the single and combined effects of phosphorus (P) availability and silver, a re-emerging contaminant, on the elemental [carbon (C):nitrogen (N):P ratios] and biochemical (fatty acid profiles) compositions of a diatom-dominated biofilm initially collected in a shallow lake. We hypothesized that (1) P and silver, through the replacement of diatoms by more tolerant primary producer species, reduce the biochemical quality of biofilms for their consumers while (2) P enhances biofilm elemental quality and (3) silver contamination of biofilm has negative effects on consumers life history traits. The quality of biofilms for consumers was assessed for a common crustacean species, Gammarus fossarum, by measuring organisms' survival and growth rates during a 42-days feeding experiment. Results mainly showed that species replacement induced by both stressors affected biofilm fatty acid compositions, and that P immobilization permitted to achieve low C:P biofilms, whatever the level of silver contamination. Gammarids growth and survival rates were not significantly impacted by the ingestion of silver-contaminated resource. On the contrary, we found a significant positive relationship between the biofilm P-content and gammarids growth. This study underlines the large indirect consequences stressors could play on the quality of microbial biomass for consumers, and, in turn, on the whole food web.

Global patterns and drivers of ecosystem functioning in rivers and riparian zones.

River ecosystems receive and process vast quantities of terrestrial organic carbon, the fate of which depends strongly on microbial activity. Variation in and controls of processing rates, however, are poorly characterized at the global scale. In response, we used a peer-sourced research network and a highly standardized carbon processing assay to conduct a global-scale field experiment in greater than 1000 river and riparian sites. We found that Earth's biomes have distinct carbon processing signatures. Slow processing is evident across latitudes, whereas rapid rates are restricted to lower latitudes. Both the mean rate and variability decline with latitude, suggesting temperature constraints toward the poles and greater roles for other environmental drivers (e.g., nutrient loading) toward the equator. These results and data set the stage for unprecedented "next-generation biomonitoring" by establishing baselines to help quantify environmental impacts to the functioning of ecosystems at a global scale.

Nutrient effects on the genetic and functional diversity of aquatic bacterial communities.

Studies on the effects of biodiversity on ecosystem functioning have generally revealed a positive asymptotic relationship between biodiversity and single functions, suggesting species redundancy with respect to these functions. However, most research was performed on specific processes and did not consider ecosystem 'multifunctionality'. There is also little information on the relationship between genetic and functional diversity. To analyze this relationship, we performed a microcosm experiment on a complex lake assemblage of decomposers, in the presence of the green alga Scenedesmus obliquus, which acted as carbon source for decomposers. By manipulating nutrient enrichment and the N : P input ratio, we observed that the structures of particle-associated and free bacterial assemblages were highly predictable in response to stoichiometric constraints. For a given treatment, the taxonomic compositions of free and particle-associated bacterial communities appeared close to each other only when phosphorus was not depleted. A coinertia analysis revealed a clear coupling between the genetic diversity of the microbial community, assessed using PCR-denatured gradient gel electrophoresis, and its potential functional diversity, studied with Biolog Ecoplates. This suggests that an ecologically relevant fraction of bacterial communities is characterized by lower level of redundancy than frequently thought, highlighting the necessity of exploring further the role of biodiversity in multifunctionality within ecosystems.

High stoichiometric food quality increases moulting organism vulnerability to pollutant impacts: An experimental test with Gammarus fossarum (Crustacea: Amphipoda).

Headwater organisms are most often simultaneously faced with multiple stressors such as low resource quality and pollutants. Higher food quality has been hypothesized to enhance the tolerance of organisms to pollutants, but the interactive effects of food quality and pollutants on species and ecosystems remain poorly studied. To better understand these interactive effects, we experimentally manipulated the phosphorus (P) content of two leaf litters with contrasted carbon quality (alder and maple). During four weeks, individuals of the detritivorous crustacean Gammarus fossarum were exposed to low levels of cadmium ([Cd] = 0, 0.35 or 0.70 µg L-1) while being fed with one of the leaf P treatments. When organisms were not exposed to Cd, their high survival rate was more driven by the carbon quality of the resource (litter species) than by its stoichiometric quality. In contrast, their number of moults and growth rates were primarily increased by the P content of resources. When exposed to Cd, G. fossarum survival rate was reduced, but this effect was largely magnified by a higher P level in resources. Our results showed that despite positive effects of resource stoichiometric quality on organism life history traits (growth, survival), a resource of high stoichiometric quality might be detrimental for organisms exposed to low and environmentally realistic levels of pollutants. Two non-exclusive hypotheses are proposed to explain these results. First, organisms fed on the highest quality resource exhibited the highest moulting frequencies (moults being the most critical life cycle step of arthropods), which could have rendered them more sensitive to pollutants. Secondly, the metabolism of organisms fed on higher quality resources was potentially enhanced, increasing the uptake of dissolved Cd by gammarids. This study suggests that species sensitivity to pollutants might be underestimated in ecosystems facing both nutrient constraint and pollutants.

Additive effect of calcium depletion and low resource quality on Gammarus fossarum (Crustacea, Amphipoda) life history traits.

Gammarus fossarum is an often-abundant crustacean detritivore that contributes importantly to leaf litter breakdown in oligotrophic, mainly heterotrophic, headwater streams. This species requires large amounts of Ca to moult, thus allowing growth and reproduction. Because resource quality is tightly coupled to the organism's growth and physiological status, we hypothesised that low Ca concentration [Ca] and low food resource quality (low phosphorus [P] and/or reduced highly unsaturated fatty acid [HUFA] contents) would interactively impair molecular responses (gene expression) and reproduction of G. fossarum. To investigate the effects of food resources quality, we experimentally manipulated the P content of sycamore leaves and also used diatoms because they contain high amounts of HUFAs. Three resource quality treatments were tested: low quality (LQ, unmanipulated leaves: low P content), high quality 1 (HQ1; P-manipulated leaves: high P content), and high quality 2 (unmanipulated leaves supplemented with a pellet containing diatoms: high P and HUFA content). Naturally, demineralised stream water was supplemented with CaSO4 to obtain three Ca concentrations (2, 3.5, and 10.5 mg Ca L-1). For 21 days, pairs of G. fossarum were individually exposed to one of the nine treatments (3 [Ca] × 3 resource qualities). At the individual level, strong and significant delays in moult stage were observed in gammarids exposed to lower [Ca] and to lower resource quality, with additive effects lengthening the duration of the reproductive cycle. Effects at the molecular level were investigated by measuring expression of 12 genes involved in energy production, translation, or Ca or P homeostasis. Expression of ATP synthase beta (higher in HQ2), calcified cuticle protein (higher in HQ1 and HQ2), and tropomyosin (higher in HQ2 compared to HQ1) was significantly affected by resource quality, and significant additive effects on Ca transporting ATPase expression were induced by [Ca] and resource quality (higher for low [Ca] and higher resource quality). These results highlight the potential drastic deleterious effects of water [Ca] depletion on G. fossarum physiology, populations, and ecosystem functioning, especially in oligotrophic environments.