Storm impacts on alpine lakes: Antecedent weather conditions matter more than the event intensity.

Extreme weather events may be just as important as gradual trends for the long-term trajectories of ecosystems. For alpine lakes, which are exposed to both exacerbated atmospheric warming and intense episodic weather events, future conditions might not be appropriately forecast by only climate change trends, i.e. warming, if extreme events have the potential to deflect their thermal and metabolic states from their seasonal ranges. We used high-frequency monitoring data over three open-water seasons with a one-dimensional hydrodynamic model of the high-altitude Lake Muzelle (France) to show that rainstorms or windstorms, notwithstanding their intensity, did not trigger long-lasting consequences to the lake characteristics when light penetration into the lake was not modified. In contrast, storms associated with high turbidity input from the watershed ("turbid storms") strongly modified the lacustrine hydrodynamics and metabolism for the rest of the open-water season through reduced light penetration. The long-lasting effects of turbid storms were related to the inputs and in-lake persistence of very light glacial suspensoids from the watershed. The occurrence of the observed turbid storms was not related to the wind or rain intensities during the events. Instead, the turbid storms occurred after dry and atypically warm spells, i.e. meteorological conditions expected to be more frequent in this alpine region in the upcoming decades. Consequently, storm events, notwithstanding their intensity, are expected to strongly imprint the future ecological status of alpine lakes under climate warming.

Tracking a century of changes in microbial eukaryotic diversity in lakes driven by nutrient enrichment and climate warming.

High-throughput sequencing of sedimentary DNA (sed-DNA) was utilized to reconstruct the temporal dynamics of microbial eukaryotic communities (MECs) at a centennial scale in two re-oligotrophicated lakes that were exposed to different levels of phosphorus enrichment. The temporal changes within the MECs were expressed in terms of richness, composition and community structure to investigate their relationships with two key forcing factors (i.e., nutrient enrichment and climate warming). Various groups, including Apicomplexa, Cercozoa, Chrysophyceae, Ciliophora, Chlorophyceae and Dinophyceae, responded to phosphorus enrichment levels with either positive or negative impacts on their richness and relative abundance. For both lakes, statistical modelling demonstrated that phosphorus concentration ([P]) was a dominant contributor to MECs modifications before the 1980s; after the mid-80s, the contribution of air temperature changes increased and potentially surpassed the contribution of [P]. Co-occurrence network analysis revealed that some clusters of taxa (i.e., modules) composed mainly of Dinophyceae and unclassified Alveolata were strongly correlated to air temperature in both lakes. Overall, our data showed that sed-DNA constitutes a precious archive of information on past biodiversity changes, allowing the study of the dynamics of numerous eukaryotic groups that were not traditionally considered in paleo-reconstructions.

Global spread of hypoxia in freshwater ecosystems during the last three centuries is caused by rising local human pressure.

The spread of hypoxia is a threat to aquatic ecosystem functions and services as well as to biodiversity. However, sparse long-term monitoring of lake ecosystems has prevented reconstruction of global hypoxia dynamics while inhibiting investigations into its causes and assessing the resilience capacity of these systems. This study compiles the onset and duration of hypoxia recorded in sediments of 365 lakes worldwide since AD 1700, showing that lacustrine hypoxia started spreading before AD 1900, 70 years prior to hypoxia in coastal zones. This study also shows that the increase of human activities and nutrient release is leading to hypoxia onset. No correlations were found with changes in precipitation or temperature. There is no evidence for a post-1980s return to well-oxygenated lacustrine conditions in industrialized countries despite the implementation of restoration programs. The apparent establishment of stable hypoxic conditions prior to AD 1900 highlights the challenges of a growing nutrient demand, accompanied by increasing global nutrient emissions of our industrialized societies, and climate change.

Particle-Dissolved Phase Partition of Polychlorinated Biphenyls in High Altitude Alpine Lakes.

We investigated whether polychlorinated biphenyls (PCBs) partitioning between the dissolved and particulate phases in two high altitude alpine lakes was determined by the quantity, size structure, or composition of suspended particles. Within- and between-lakes differences in water-particulate phase partition coefficient (Kp) were not related to total suspended matter, phytoplankton biomass, or taxonomic composition. Yet, a seasonal relationship between Kp and Kow was detected for both lakes, revealing equilibrium of PCBs partition when lakes were ice covered. On the contrary, PCBs partitioning between particles and water appeared kinetically limited during the open water season. Partition is therefore mainly governed by thermodynamic laws during the ice-covered period, while none of the tested physical or biological parameters seemed to explain the distribution of these particle-reactive contaminants in the open water period. PCBs were always mainly associated with particulate matter, but partitioning within different particulate size-fractions varied between seasons and between years during open water periods. When ice cover is absent, PCBs were mainly adsorbed on microplankton, the largest phytoplanktonic size fraction, which is the least likely to get grazed by pelagic microconsumers.

Historical profiles of PCB in dated sediment cores suggest recent lake contamination through the "halo effect".

We investigated the major sources of polychlorinated biphenyls (PCB) and interpreted the environmental fate processes of these persistent organic pollutants in the past and current PCB contamination of three large, urbanized, French peri-alpine lakes. Dated sediment cores were analyzed in order to reconstruct and compare the historical contamination in all three lakes. Stratigraphic changes of PCB contents and fluxes were considered as revealing the temporal dynamics of PCB deposition to the lakes and the distribution of the seven indicator congeners (further referred to as PCBi) as an indicator of the main contamination origin and pathway. Although located within a single PCB industrial production region, concentration profiles for the three lakes differed in timing, peak concentration magnitudes, and in the PCBi congeners compositions. PCBi fluxes to the sediment and the magnitude of the temporal changes were generally much lower in Lake Annecy (0.05-2 ng·cm(-2)·yr(-1)) as compared to Lakes Geneva (0.05-5 ng·cm(-2)·yr(-1)) and Bourget (5-290 ng·cm(-2)·yr(-1)). For all three lakes, the paramount contamination occurred in the early 1970s. In Lakes Annecy and Bourget, PCB fluxes have declined and plateaued at 0.5 and 8 ng·cm(-2)·yr(-1), respectively, since the early 1990s. In Lake Geneva, PCB fluxes have further decreased by the end of the XX(th) century and are now very low. For the most contaminated lake (Lake Bourget), the high PCBi flux (5-290 ng·cm(-2)·yr(-1)) and the predominance of heavy congeners for most of the time period are consistent with a huge local input to the lake. This still high rate of Lake Bourget is explained by transport of suspended solids from one of its affluents, polluted by an industrial point source. Intermediate historical levels and PCBi distribution over time for Lake Geneva suggest a mixed contamination (urban point sources and distant atmospheric transport), while atmospheric deposition to Lake Annecy explains its lowest contamination rate. The presently low contamination levels recorded in Lake Geneva correspond to atmospheric inputs, but the recent PCBi distribution of Lake Annecy, enriched in relatively heavy congeners, reveals a contamination by the neighboring Lake Bourget, following a halo effect of about 40 km radius.

Changes in carbon sources fueling benthic secondary production over depth and time: coupling Chironomidae stable carbon isotopes to larval abundance.

Stable C isotope ratio (δ(13)C) values of chironomid remains (head capsules; HC) were used to infer changes in benthic C sources over the last 150 years for two French sub-Alpine lakes. The HCs were retrieved from a series of sediment cores from different depths. The HC δ(13)C values started to decrease with the onset of eutrophication. The HC δ(13)C temporal patterns varied among depths, which revealed spatial differences in the contribution of methanotrophic bacteria to the benthic secondary production. The estimates of the methane (CH4)-derived C contribution to chironomid biomass ranged from a few percent prior to the 1930s to up to 30 % in recent times. The chironomid fluxes increased concomitantly with changes in HC δ(13)C values before a drastic decrease due to the development of hypoxic conditions. The hypoxia reinforced the implication for CH4-derived C transfer to chironomid production. In Lake Annecy, the HC δ(13)C values were negatively correlated to total organic C (TOC) content in the sediment (Corg), whereas no relationship was found in Lake Bourget. In Lake Bourget, chironomid abundances reached their maximum with TOC contents between 1 and 1.5 % Corg, which could constitute a threshold for change in chironomid abundance and consequently for the integration of CH4-derived C into the lake food webs. Our results indicated that the CH4-derived C contribution to the benthic food webs occurred at different depths in these two large, deep lakes (deep waters and sublittoral zone), and that the trophic transfer of this C was promoted in sublittoral zones where O2 gradients were dynamic.

Lake surface cooling drives littoral-pelagic exchange of dissolved gases.

The extent of littoral influence on lake gas dynamics remains debated in the aquatic science community due to the lack of direct quantification of lateral gas transport. The prevalent assumption of diffusive horizontal transport in gas budgets fails to explain anomalies observed in pelagic gas concentrations. Here, we demonstrate through high-frequency measurements in a eutrophic lake that daily convective horizontal circulation generates littoral-pelagic advective gas fluxes one order of magnitude larger than typical horizontal fluxes used in gas budgets. These lateral fluxes are sufficient to redistribute gases at the basin-scale and generate concentration anomalies reported in other lakes. Our observations also contrast the hypothesis of pure, nocturnal littoral-to-pelagic exchange by showing that convective circulation transports gases such as oxygen and methane toward both the pelagic and littoral zones during the daytime. This study challenges the traditional pelagic-centered models of aquatic systems by showing that convective circulation represents a fundamental lateral transport mechanism to be integrated into gas budgets.

Local forcings affect lake zooplankton vulnerability and response to climate warming.

While considerable insights on the ecological consequences of climate change have been gained from studies conducted on remote lakes, little has been done on lakes under direct human exposure. Ecosystem vulnerability and responses to climate warming might yet largely depend on the ecological state and thus on local anthropogenic pressures. We tested this hypothesis through a paleolimnological approach on three temperate large lakes submitted to rather similar climate warming but varying intensities of analogous local forcings (changes in nutrient inputs and fisheries management practices). Changes in the structure of the cladoceran community were considered as revealing for alterations, over the time, of the pelagic food web. Trajectories of the cladoceran communities were compared among the three study lakes (Lakes Geneva, Bourget, and Annecy) over the last 70-150 years. Generalized additive models were used to develop a hierarchical understanding of the respective roles of local stressors and climate warming in structuring cladoceran communities. The cladoceran communities were not equally affected by climate warming between lakes. In Lake Annecy, which is the most nutrient-limited, the cladoceran community was essentially controlled by local stressors, with very limited impact of climate. In contrast, the more nutrient-loaded Lakes Geneva and Bourget were more sensitive to climate warming, although the magnitude of their responses and the pathways under which climate warming affected the communities varied between the two lakes. Finally, our results demonstrated that lake vulnerability and responses to climate warming are modulated by lake trophic status but can also be altered by fisheries management practices through changes in fish predation pressure.

Are cyanobacterial blooms trophic dead ends?

Cyanobacterial blooms induce significant costs that are expected to increase in the near future. Cyanobacterial resistance to zooplankton grazing is one factor thought to promote bloom events. Yet, numerous studies on zooplankton ability to graze upon cyanobacteria have been producing contradictory results and such a puzzle might arise from the lack of direct observations in situ. Our objective was to track, using fatty acid (FA) and fatty acid stable isotope analyses (FA-SIA), the fate of cyanobacterial organic matter in the food web of a lake subjected to summer blooms of Planktothrix rubescens. A metalimnetic bloom of P. rubescens occurred in Lake Bourget (France) during the study period (May-November 2009). The bloom was especially rich in α-linolenic acid, 18:3(n-3), but none of the considered zooplankton taxa exhibited spiking content in this particular FA. FA-SIA revealed, however, that over a quarter of 18:3(n-3) in small zooplankton (<500 µm) was provided by P. rubescens while large cladocerans (>500 µm) did not benefit from it. P. rubescens 18:3(n-3) could be tracked up to perch (Perca fluviatilis) young of the year (YOY) to which it contributed to ~15 % of total 18:3(n-3). Although transferred with a much lower efficiency than micro-algal organic matter, the P. rubescens bloom supported a significant share of the pelagic secondary production and did not constitute, sensu stricto, a 'trophic dead end'. The cyanobacterial bloom also provided perch YOY with components of high nutritional values at a season when these are critical for their recruitment. This cyanobacterial bloom might thus be regarded as a significant dietary bonus for juvenile fish.

Fine-scale dynamics of calcite precipitation in a large hardwater lake.

In hardwater lakes, calcite precipitation is an important yet poorly understood process in the lacustrine carbon cycle, in which catchment-derived alkalinity (Alk) is both transformed and translocated. While the physico-chemical conditions supporting the supersaturation of water with respect to calcite are theoretically well described, the magnitude and conditions underlying calcite precipitation at fine temporal and spatial scales are poorly constrained. In this study, we used high frequency, depth-resolved (0-30 m) data collected over 18 months (June 2019 - November 2020) in the deeper basin of Lake Geneva to describe the dynamics of calcite precipitation fluxes at a fine temporal resolution (day to season) and to scale them to carbon fixation by primary production. Calcite precipitation occurred during the warm stratified periods when surface water CO2 concentrations were below atmospheric equilibrium. Seasonally, the extent of Alk loss due to calcite precipitation (i.e., [30-42] g C m-2) depended upon the level of Alk in surface waters. Moreover, interannual variability in seasonal calcite precipitation depended on the duration of stratification, which determined the volume of the water layer susceptible to calcite precipitation. At finer timescales, calcite precipitation was characterized by marked daily variability with dynamics strongly related to that of planktonic autotrophic metabolism. Increasing daily calcite precipitation rates (i.e., maximum values 9 mmol C m-3 d-1) coincided with increasing net ecosystem production (NEP) during periods of enhanced water column stability. In these conditions, calcite precipitation could remove as much inorganic carbon from the productive layers as NEP. This study provides mechanistic insights into the conditions driving pelagic calcite precipitation, and quantifies its essential contribution to the coupling of organic and inorganic carbon cycling in lakes.

Intense touristic activities exceed climate change to shape aquatic communities in a mountain lake.

Mountain lakes are especially vulnerable to climate change, but are also increasingly exposed to local anthropogenic development through winter and summer tourism. In this study, we aimed to tease apart the influence of tourism from that of climate in a mountain lake located within one of the largest French ski resorts, by combining paleolimnological and present ecological data. The reconstructed long-term ecological dynamics highlighted an increase in lake biological production from the end of the Little Ice Age up to the 1950s, suggesting a historical dominance of climate control. Afterward, a major drop in pelagic production occurred at the same time as the watershed erosion increased and peaked in the 1990s, concomitant with massive digging for the ski resort expansion. The benthic invertebrates collapsed in the 1980s, concomitantly with the onset of massive salmonid stocking and recent warming. Stable isotope analyses identified benthic invertebrates as the major salmonid diet resource and suggested a possible direct impact of salmonid stocking on benthic invertebrates. However, habitat use may differ among salmonid species as suggested by the way fish DNA was preserved in surficial sediment. The high abundances of macrozooplankton further confirmed the limited reliance of salmonids on pelagic resources. The variable thermal tolerance of benthic invertebrates suggested that the recent warming may mostly affect littoral habitats. Our results indicate that winter and summer tourism may differently affect the biodiversity of mountain lakes and could collectively interfere with the ecological impacts of recent warming, making local management of primary importance to preserve their ecological integrity. Supplementary Information: The online version contains supplementary material available at 10.1007/s00027-023-00968-6.

Quantitative PCR enumeration of total/toxic Planktothrix rubescens and total cyanobacteria in preserved DNA isolated from lake sediments.

The variability of spatial distribution and the determinism of cyanobacterial blooms, as well as their impact at the lake scale, are still not understood, partly due to the lack of long-term climatic and environmental monitoring data. The paucity of these data can be alleviated by the use of proxy data from high-resolution sampling of sediments. Coupling paleolimnological and molecular tools and using biomarkers such as preserved DNA are promising approaches, although they have not been performed often enough so far. In our study, a quantitative PCR (qPCR) technique was applied to enumerate total cyanobacterial and total and toxic Planktothrix communities in preserved DNA derived from sediments of three lakes located in the French Alps (Lake Geneva, Lake Bourget, and Lake Annecy), containing a wide range of cyanobacterial species. Preserved DNA from lake sediments was analyzed to assess its quality, quantity, and integrity, with further application for qPCR. We applied the qPCR assay to enumerate the total cyanobacterial community, and multiplex qPCR assays were applied to quantify total and microcystin-producing Planktothrix populations in a single reaction tube. These methods were optimized, calibrated, and applied to sediment samples, and the specificity and reproducibility of qPCR enumeration were tested. Accurate estimation of potential inhibition within sediment samples was performed to assess the sensitivity of such enumeration by qPCR. Some precautions needed for interpreting qPCR results in the context of paleolimnological approaches are discussed. We concluded that the qPCR assay can be used successfully for the analysis of lake sediments when DNA is well preserved in order to assess the presence and dominance of cyanobacterial and Planktothrix communities.

Parasitic versus nutritional regulation of natural fish populations.

Although parasites are expected to affect their host's fitness, quantitative proof for impacts of parasitism on wild populations is hampered by confounding environmental factors, including dietary resource. Herein, we evaluate whether the physiological conditions of European perch (Perca fluviatilis) in three large peri-alpine lakes (Geneva, Annecy, and Bourget) depend on (a) the nutritional status of the juvenile fish, as revealed by stable isotope and fatty acid compositions, (b) the prevalence of the tapeworm Triaenophorus nodulosus, a parasite transmitted to perch through copepod preys, or (c) interactive effects of both factors. At the scale of lake populations, the deficit in growth and fat storage of juvenile perch during their first summer coincides with a high parasite prevalence and also a low quality of dietary resource. Yet, at the individual level, parasites had no evident effect on the growth of the juvenile perch, while impacts on fat storage appeared only at the highest prevalence of the most infected lake. Fatty acid and stable isotope analyses of fish tissue do not reveal any impact of T. nodulosus on diet, physiology, and feeding behaviour of fish within lakes. Overall, we found a low impact of parasitism on the physiological condition and trophic status of juvenile perch at the end of their first summer. We find instead that juvenile perch growth and fat storage, both factors tied to their winter survival, are under strong nutritional constraints. However, the coinciding nutritional constraints and parasite prevalence of perch juveniles in these three lakes may result from the indirect effect of lake nutrient concentrations, which, as a major control of zooplankton communities, simultaneously regulate both the dietary quality of fish prey and the host-parasite encounter rates.

[Strengths, weaknesses, and opportunities of French research in trophic ecology]. / Forces, failles et opportunités de la recherche française en écologie trophique.

The French National Institute of Ecology and Environment (INEE) aims at fostering pluridisciplinarity in Environmental Science and, for that purpose, funds ex muros research groups (GDR) on thematic topics. Trophic ecology has been identified as a scientific field in ecology that would greatly benefit from such networking activity, as being profoundly scattered. This has motivated the seeding of a GDR, entitled "GRET". The contours of the GRET's action, and its ability to fill these gaps within trophic ecology at the French national scale, will depend on the causes of this relative scattering. This study relied on a nationally broadcasted poll aiming at characterizing the field of trophic ecology in France. Amongst all the unique individuals that fulfilled the poll, over 300 belonged at least partly to the field of trophic ecology. The sample included all French public research institutes and career stages. Three main disruptions within the community of scientist in trophic ecology were identified. The first highlighted the lack of interfaces between microbial and trophic ecology. The second evidenced that research questions were strongly linked to single study fields or ecosystem type. Last, research activities are still quite restricted to the ecosystem boundaries. All three rupture points limit the conceptual and applied progression in the field of trophic ecology. Here we show that most of the disruptions within French Trophic Ecology are culturally inherited, rather than motivated by scientific reasons or justified by socio-economic stakes. Comparison with the current literature confirms that these disruptions are not necessarily typical of the French research landscape, but instead echo the general weaknesses of the international research in ecology. Thereby, communication and networking actions within and toward the community of trophic ecologists, as planned within the GRET's objectives, should contribute to fill these gaps, by reintegrating microbes within trophic concepts and setting the seeds for trans- and meta-ecosystemic research opportunities. Once the community of trophic ecologists is aware of the scientific benefit in pushing its boundaries forwards, turning words and good intentions into concrete research projects will depend on the opportunities to obtain research funding.

Seasonal variations in fish delta13C and delta15N in two West African reservoirs, Sélingué and Manantali (Mali): modifications of trophic links in relation to water level.

Most regions in the tropics undergo high seasonal precipitation that produces cyclic patterns of riverine discharge, resulting in periods characterized by low and high water levels. Many chemical and bio-logical factors are affected by this hydrologic seasonality, and it therefore appeared to be very likely that aquatic food webs would also differ during the low and high water periods. Available carbon sources for fish are thought to be less varied during low water periods, but flooding during high water periods could bring fish into contact with a greater abundance and diversity of food sources such as terrestrial plants or the biofilms that grow on submerged terrestrial plants. At low water levels, higher fish densities may lead to more piscivory and less omnivory when compared with the high water periods. Therefore, trophic links within the fish communities may then be modified by water level changes in tropical reservoirs. To address this prediction, we performed stable isotope analyses of the most common species in Sélingué and Manantali, two large reservoirs in Mali (West Africa). Allochthonous and littoral carbon sources were shown to support fish production to a significant extent, even during low water periods. However, the allochthonous or littoral carbon contributions that sustained the top-predators production were indeed greater during the high water periods as expected. The expected higher omnivory in the high water period might have shortened the food chain when compared with the low water period. Some carnivorous fish species were shown to feed at lower trophic levels during high water periods in both reservoirs, but this was not a general pattern. Flooding did not, therefore, necessarily result in a shorter food chain when water levels were high.

Mass budget in two high altitude lakes reveals their role as atmospheric PCB sinks.

A mass budget of polychlorinated biphenyls (PCBs) was constructed for two altitude lakes located in the French Alps to (i) quantify inward and outward PCB flux over the entire year of 2012, (ii) hierarchize the dominant pathways of PCB transfers, and (iii) evaluate to what extent these pathways vary between both lakes. The annual PCB inputs were similar, and the glacial runoff and sediment-to-water exchange were negligible sources of PCBs to the water column relative to atmospheric deposition. The annual inputs were primarily introduced by snow deposition and transferred into the lakes during the few weeks of spring thaw. While the dominant deposition pathways were similar, the main processes by which the water column lost pollutants differed between the two lakes. Despite these differences, the mass budget revealed that PCB inputs exceeded outputs for both studied lakes and that the lakes acted as atmospheric PCB sinks for the surrounding mountain environment. The differences in the PCB distribution between the key compartments (sediment and water column) are most likely due to differences in the lacustrine internal processes.