The chemical succession in anoxic lake waters as source of molecular diversity of organic matter.

The aquatic networks that connect soils with oceans receive each year 5.1 Pg of terrestrial carbon to transport, bury and process. Stagnant sections of aquatic networks often become anoxic. Mineral surfaces attract specific components of organic carbon, which are released under anoxic conditions to the pool of dissolved organic matter (DOM). The impact of the anoxic release on DOM molecular composition and reactivity in inland waters is unknown. Here, we report concurrent release of iron and DOM in anoxic bottom waters of northern lakes, removing DOM from the protection of iron oxides and remobilizing previously buried carbon to the water column. The deprotected DOM appears to be highly reactive, terrestrially derived and molecularly distinct, generating an ambient DOM pool that relieves energetic constraints that are often assumed to limit carbon turnover in anoxic waters. The Fe-to-C stoichiometry during anoxic mobilization differs from that after oxic precipitation, suggesting that up to 21% of buried OM escapes a lake-internal release-precipitation cycle, and can instead be exported downstream. Although anoxic habitats are transient and comprise relatively small volumes of water on the landscape scale, our results show that they may play a major role in structuring the reactivity and molecular composition of DOM transiting through aquatic networks and reaching the oceans.

Integrating terrestrial and aquatic ecosystems to constrain estimates of land-atmosphere carbon exchange.

In this Perspective, we put forward an integrative framework to improve estimates of land-atmosphere carbon exchange based on the accumulation of carbon in the landscape as constrained by its lateral export through rivers. The framework uses the watershed as the fundamental spatial unit and integrates all terrestrial and aquatic ecosystems as well as their hydrologic carbon exchanges. Application of the framework should help bridge the existing gap between land and atmosphere-based approaches and offers a platform to increase communication and synergy among the terrestrial, aquatic, and atmospheric research communities that is paramount to advance landscape carbon budget assessments.

Catchment-scale carbon fluxes and processes in major rivers of northern Québec, Canada.

Boreal rivers transport and process large amounts of organic and inorganic materials derived from their catchments, yet quantitative estimates and patterns of carbon (C) transport and emissions in these large rivers are scarce relative to those of high-latitude lakes and headwater streams. Here, we present the results of a large-scale survey of 23 major rivers in northern Québec sampled during the summer period of 2010, which aimed to determine the magnitude and spatial variability of different C species (carbon dioxide - CO2, methane - CH4, total carbon - TC, dissolved organic carbon - DOC and inorganic carbon - DIC), as well as to identify their main drivers. In addition, we constructed a first order mass balance of total riverine C emissions to the atmosphere (outgassing from the main river channel) and export to the ocean over summer. All rivers were supersaturated in pCO2 and pCH4 (partial pressure of CO2 and CH4), and the resulting fluxes varied widely among rivers, especially the CH4. There was a positive relationship between DOC and gas concentrations, suggesting a common watershed source of these C species. DOC concentrations declined as a function of % land surface covered by water (lentic + lotic systems) in the watershed, suggesting that lentic systems may act as a net sink of organic matter in the landscape. The C balance suggests that the export component is higher than atmospheric C emissions in the river channel. However, for heavily dammed rivers, C emissions to the atmosphere approaches the C export component. Such studies are highly important for the overall efforts to effectively quantify and incorporate major boreal rivers into whole-landscape C budgets, to determine the net role of these ecosystems as C sinks or sources, and to predict how these might shift under anthropogenic pressures and dynamic climate conditions.

Terrestrial connectivity, upstream aquatic history and seasonality shape bacterial community assembly within a large boreal aquatic network.

During transit from soils to the ocean, microbial communities are modified and re-assembled, generating complex patterns of ecological succession. The potential effect of upstream assembly on downstream microbial community composition is seldom considered within aquatic networks. Here, we reconstructed the microbial succession along a land-freshwater-estuary continuum within La Romaine river watershed in Northeastern Canada. We captured hydrological seasonality and differentiated the total and reactive community by sequencing both 16 S rRNA genes and transcripts. By examining how DNA- and RNA-based assemblages diverge and converge along the continuum, we inferred temporal shifts in the relative importance of assembly processes, with mass effects dominant in spring, and species selection becoming stronger in summer. The location of strongest selection within the network differed between seasons, suggesting that selection hotspots shift depending on hydrological conditions. The unreactive fraction (no/minor RNA contribution) was composed of taxa with diverse potential origins along the whole aquatic network, while the majority of the reactive pool (major RNA contribution) could be traced to soil/soilwater-derived taxa, which were distributed along the entire rank-abundance curve. Overall, our findings highlight the importance of considering upstream history, hydrological seasonality and the reactive microbial fraction to fully understand microbial community assembly on a network scale.

Modeling the spatial and temporal variability in surface water CO2 and CH4 concentrations in a newly created complex of boreal hydroelectric reservoirs.

Hydroelectric reservoirs emit carbon dioxide (CO2) and methane (CH4) to the atmosphere, yet there is still much uncertainty concerning the magnitude and drivers of these greenhouse gas (GHG) emissions. This uncertainty is particularly large over the initial years after flooding and in complex, cascade reservoir systems where studies are rare. We assessed the spatial and temporal patterns of CO2 and CH4 concentrations in the newly created La Romaine complex, which is composed of three consecutive reservoirs (RO1, RO2, RO3) along the La Romaine River. Dissolved CO2 and CH4 concentrations were intensively measured over three seasons for four years. Results show elevated CH4 and especially CO2 concentrations in surface waters of all three reservoirs upon flooding, with strong seasonality and high spatial heterogeneity within reservoirs. There was a strong seasonal decoupling of surface water CO2 and CH4 concentrations. Contrary to expectations, surface water CO2 and CH4 concentrations were relatively stable over the initial years of flooding, with exception of the decrease in CO2 concentrations in the shallower RO1 reservoir. Further, individual reservoir characteristics, notably reservoir morphometry and pre-flood land cover, together with climatic factors were the main drivers of CO2 and CH4 concentrations, and the reservoir position in the cascade played a minor role. Models differed for CO2 and CH4, and also between reservoirs highlighting the need to capture these specificities in reservoir functioning. We establish a modeling framework to effectively fill the spatial and temporal gaps that inevitably exist in the sampling coverage of large and heterogeneous reservoirs, which combined with appropriately modeled gas transfer velocities, will serve as a platform to derive robust estimates of diffusive fluxes. This modeling framework can be transposed to other reservoirs, and will contribute to more accurate and representative estimates of diffusive carbon emissions from hydroelectric reservoirs.

Tracking the upstream history of aquatic microbes in a boreal lake yields new insights on microbial community assembly.

Bacterial community structure can change rapidly across short spatial and temporal scales as environmental conditions vary, but the mechanisms underlying those changes are still poorly understood. Here, we assessed how a lake microbial community assembles by following its reorganization from the main tributary, which, when flowing into the lake, first traverses an extensive macrophyte-dominated vegetated habitat, before reaching the open water. Environmental conditions in the vegetated habitat changed drastically compared to both river and lake waters and represented a strong environmental gradient for the incoming bacteria. We used amplicon sequencing of the 16S rRNA gene and transcript to reconstruct the shifts in relative abundance of individual taxa and link this to their pattern in activity (here assessed with RNA:DNA ratios). Our results indicate that major shifts in relative abundance were restricted mostly to rare taxa (<0.1% of relative abundance), which seemed more responsive to environmental changes. Dominant taxa (>1% of relative abundance), on the other hand, traversed the gradient mostly unchanged with relatively low and stable RNA:DNA ratios. We also identified a high level of local recruitment and a seedbank of taxa capable of activating/inactivating, but these were almost exclusively associated with the rare biosphere. Our results suggest a scenario where the lake community results from a reshuffling of the rank abundance structure within the incoming rare biosphere, driven by selection and growth, and that numerical dominance is not a synonym of activity, growth rate, or environmental selection, but rather reflect mass effects structuring these freshwater bacterial communities.

Freshwater zooplankton metapopulations and metacommunities respond differently to environmental and spatial variation.

Theory predicts that population genetic structure and metacommunity structure are linked by the common processes of drift and migration, but how population genetic structure and metacommunity structure are related in nature is still unknown. Deeper understanding of the processes influencing both genetic and community diversity is vital for better predicting how environmental change will impact biodiversity patterns. We examined how crustacean zooplankton and rotifer species' metapopulation genetic structure and metacommunities respond to environmental and spatial variation both within and across four regions of boreal Canada. Metapopulation and metacommunity variation partitioning results were compared within and across the four regions. Metapopulations and metacommunities responded differently to environmental variation and spatial structure both within and across regions, as metapopulations were influenced by different environmental variables compared to metacommunities. At larger spatial scales both metapopulations and metacommunities exhibited greater spatial and environmental structuring, again responding to a different subset of environmental variables. Our findings suggest that even though both genetic and species diversity are linked by the same processes, regional variation in environmental characteristics and spatial structure influence resulting biodiversity patterns differently. To date, no other empirical research has explored relationships between entire metapopulation and metacommunity assemblages at large regional spatial scales.

Comment on "On the calculation of lake metabolic rates: Diel O2 and 18/16O technique" by Peeters et al. [Water Res. 165 2019, 114990].

Quantifying metabolic rates in lakes and other aquatic ecosystems is a complex task, as methods are continually evolving and are not currently standardized. Recently, Peeters et al. presented a valuable simulated dataset that advances the field by comparing the strengths and limitations of individual and combined metabolic techniques. The authors conclude that calculating metabolic rates from point sampling and mass balancing of surface water oxygen concentration and isotope composition is flawed, because the technique does not capture sub-daily patterns of metabolic variability, which they argue invalidates past applications and interpretations. These conclusions are inconsistent with how the method has been used, and are based on a biased construction of scenarios and interpretation of model results, especially because their parameterization of the stable isotopic model employs input values that appear unrepresentative of most lake conditions. Here, we establish that 1) empirical evidence supports the isotopic approach's suitability to approximate daily or longer metabolic patterns in most lakes. 2) The authors' own simulations show agreement between metabolic estimates from point isotopic measurements and average metabolic rates under most scenarios. 3) The authors' invalidation of isotopic measurements are based on the most extreme model deviations observed in simulated hypereutrophic environments. While we welcome a critical evaluation of the isotopic approach, we argue that isotopic model uncertainty needs to be placed within an appropriate context. We emphasize that isotopic sampling and steady state metabolic modelling has a key role to play in constraining metabolic patterns in the global lake landscape, but that the research questions addressed with the method need to be commensurate with the limitations and uncertainties of the approach.

Taxonomic differences shape the responses of freshwater aerobic anoxygenic phototrophic bacterial communities to light and predation.

Aerobic anoxygenic phototrophic (AAP) bacteria are a phylogenetically diverse and ubiquitous group of prokaryotes that use organic matter but can harvest light using bacteriochlorophyll a. Although the factors regulating AAP ecology have long been investigated through field surveys, the few available experimental studies have considered AAPs as a group, thus disregarding the potential differential responses between taxonomically distinct AAP assemblages. Here, we used sequencing of the pufM gene to describe the diversity of AAPs in 10 environmentally distinct temperate lakes, and to investigate the taxonomic responses of AAP communities in these lakes when subjected to similar experimental manipulations of light and predator removal. The studied communities were clearly dominated by Limnohabitans AAP but presented a clear taxonomic segregation between lakes presumably driven by local conditions, which was maintained after experimental manipulations. Predation reduction (but not light exposure) caused significant compositional shifts across most assemblages, but the magnitude of these changes could not be clearly related to changes in bulk AAP abundances or taxonomic richness of AAP assemblages during experiments. Only a few operational taxonomic units, which differed taxonomically between lakes, were found to respond positively during experimental treatments. Our results highlight that different freshwater AAP communities respond differently to similar control mechanisms, highlighting that in-depth knowledge on AAP diversity is essential to understand the ecology and potential role of these photoheterotrophs.

Efficiency of crustacean zooplankton in transferring allochthonous carbon in a boreal lake.

Increased incorporation of terrestrial organic matter (t-OM) into consumer biomass (allochthony) is believed to reduce growth capacity. In this study, we examined the relationship between crustacean zooplankton allochthony and production in a boreal lake that displays strong seasonal variability in t-OM inputs. Contrary to our hypotheses, we found no effect of allochthony on production at the community and the species levels. The high-frequency seasonal sampling (time-for-space) allowed for estimating the efficiency of zooplankton in converting this external carbon source to growth. From the daily t-OM inputs in the lake (57-3,027 kg C/d), the zooplankton community transferred 0.2% into biomass (0.01-2.36 kg C/d); this level was of the same magnitude as the carbon transfer efficiency for algal-derived carbon (0.4%). In the context of the boundless carbon cycle, which integrates inland waters as a biologically active component of the terrestrial landscape, the use of the time-for-space approach for the quantifying of t-OM trophic transfer efficiency by zooplankton is a critical step toward a better understanding of the effects of increasing external carbon fluxes on pelagic food webs.

Magnitude and drivers of integrated fluvial network greenhouse gas emissions across the boreal landscape in Québec.

Streams and rivers are now recognized to be sites of intense carbon (C) emissions, yet the lack of C emission estimates that integrate beyond individual river systems has slowed their inclusion in landscape C budgets. Here we apply empirical models of CO2 and CH4 concentrations and gas exchange continuously along entire fluvial networks to derive the total fluvial CO2 and CH4 emissions in large (3000 to 30,000 km2) watersheds located across the boreal biome of Québec (Canada). We assess how total fluvial network C emissions vary with landscape and climate properties, and compare their magnitude to other components of the landscape C budget. The total fluvial network emissions expressed as per unit watershed area ranged from 0.7 to 29.2 g C m-2 yr-1 for CO2, and 4-1780 mg C m-2 yr-1 for CH4, and neither was related to watershed area or drainage density. Rather, watershed slope and terrestrial net productivity were major drivers of the integrated network fluvial emissions. We also show that steeper watersheds had a greater proportion of emissions relative to downstream export of C from the watershed. Integrated fluvial emissions are of the same magnitude as the terrestrial C sink, yet these two fundamental components of the boreal landscape C budget are not tightly coupled.

Reactivity, fate and functional roles of dissolved organic matter in anoxic inland waters.

The transit of organic matter (OM) through the aquatic compartment of its global cycle has been intensively studied, traditionally with a focus on the processing and degradation of its dissolved fraction (dissolved organic matter, DOM). Because this is so intimately related to oxidation, the notion tenaciously persists that where oxygen is absent, DOM turnover is markedly slowed. In this Opinion Piece, we outline how diverse processes shape, transform and degrade DOM also in anoxic aquatic environments, and we focus here on inland waters as a particular case study. A suite of biogeochemical DOM functions that have received comparatively little attention may only be expressed in anoxic conditions and may result in enhanced biogeochemical roles of these deoxygenated habitats on a network scale.

Large-scale biogeography and environmental regulation of methanotrophic bacteria across boreal inland waters.

Aerobic methanotrophic bacteria (methanotrophs) use methane as a source of carbon and energy, thereby mitigating net methane emissions from natural sources. Methanotrophs represent a widespread and phylogenetically complex guild, yet the biogeography of this functional group and the factors that explain the taxonomic structure of the methanotrophic assemblage are still poorly understood. Here, we used high-throughput sequencing of the 16S rRNA gene of the bacterial community to study the methanotrophic community composition and the environmental factors that influence their distribution and relative abundance in a wide range of freshwater habitats, including lakes, streams and rivers across the boreal landscape. Within one region, soil and soil water samples were additionally taken from the surrounding watersheds in order to cover the full terrestrial-aquatic continuum. The composition of methanotrophic communities across the boreal landscape showed only a modest degree of regional differentiation but a strong structuring along the hydrologic continuum from soil to lake communities, regardless of regions. This pattern along the hydrologic continuum was mostly explained by a clear niche differentiation between type I and type II methanotrophs along environmental gradients in pH, and methane concentrations. Our results suggest very different roles of type I and type II methanotrophs within inland waters, the latter likely having a terrestrial source and reflecting passive transport and dilution along the aquatic networks, but this is an unresolved issue that requires further investigation.

The NSERC Canadian Lake Pulse Network: A national assessment of lake health providing science for water management in a changing climate.

The distribution and quality of water resources vary dramatically across Canada, and human impacts such as land-use and climate changes are exacerbating uncertainties in water supply and security. At the national level, Canada has no enforceable standards for safe drinking water and no comprehensive water-monitoring program to provide detailed, timely reporting on the state of water resources. To provide Canada's first national assessment of lake health, the NSERC Canadian Lake Pulse Network was launched in 2016 as an academic-government research partnership. LakePulse uses traditional approaches for limnological monitoring as well as state-of-the-art methods in the fields of genomics, emerging contaminants, greenhouse gases, invasive pathogens, paleolimnology, spatial modelling, statistical analysis, and remote sensing. A coordinated sampling program of about 680 lakes together with historical archives and a geomatics analysis of over 80,000 lake watersheds are used to examine the extent to which lakes are being altered now and in the future, and how this impacts aquatic ecosystem services of societal importance. Herein we review the network context, objectives and methods.

Understanding how microbiomes influence the systems they inhabit.

Translating the ever-increasing wealth of information on microbiomes (environment, host or built environment) to advance our understanding of system-level processes is proving to be an exceptional research challenge. One reason for this challenge is that relationships between characteristics of microbiomes and the system-level processes that they influence are often evaluated in the absence of a robust conceptual framework and reported without elucidating the underlying causal mechanisms. The reliance on correlative approaches limits the potential to expand the inference of a single relationship to additional systems and advance the field. We propose that research focused on how microbiomes influence the systems they inhabit should work within a common framework and target known microbial processes that contribute to the system-level processes of interest. Here, we identify three distinct categories of microbiome characteristics (microbial processes, microbial community properties and microbial membership) and propose a framework to empirically link each of these categories to each other and the broader system-level processes that they affect. We posit that it is particularly important to distinguish microbial community properties that can be predicted using constituent taxa (community-aggregated traits) from those properties that cannot currently be predicted using constituent taxa (emergent properties). Existing methods in microbial ecology can be applied to more explicitly elucidate properties within each of these three categories of microbial characteristics and connect them with each other. We view this proposed framework, gleaned from a breadth of research on environmental microbiomes and ecosystem processes, as a promising pathway with the potential to advance discovery and understanding across a broad range of microbiome science.

Soils associated to different tree communities do not elicit predictable responses in lake bacterial community structure and function.

Freshwater bacterioplankton communities are influenced by the inputs of material and bacteria from the surrounding landscape, yet few studies have investigated how different terrestrial inputs affect bacterioplankton. We examined whether the addition of soils collected under various tree species combinations differentially influences lake bacterial communities. Lake water was incubated for 6 days following addition of five different soils. We assessed the taxonomic composition (16S rRNA gene sequencing) and metabolic activity (Biolog Ecoplates) of lake bacteria with and without soil addition, and compared these to initial soil communities. Soil bacterial assemblages showed a strong influence of tree composition, but such community differences were not reflected in the structure of lake communities that developed during the experiment. Bacterial taxa showing the largest abundance increases during incubation were initially present in both lake water and across most soils, and were related to Cytophagales, Burkholderiales and Rhizobiales. No clear metabolic profiles based on inoculum source were found, yet soil-amended communities used 60% more substrate than non-inoculated communities. Overall, we show that terrestrial inputs influence aquatic communities by stimulating the growth and activity of certain ubiquitous taxa distributed across the terrestrial-aquatic continuum, yet different forest soils did not cause predictable changes in lake bacterioplankton assemblages.

Modelling CO2 emissions from water surface of a boreal hydroelectric reservoir.

To quantify CO2 emissions from water surface of a reservoir that was shaped by flooding the boreal landscape, we developed a daily time-step reservoir biogeochemistry model. We calibrated the model using the measured concentrations of dissolved organic and inorganic carbon (C) in a young boreal hydroelectric reservoir, Eastmain-1 (EM-1), in northern Quebec, Canada. We validated the model against observed CO2 fluxes from an eddy covariance tower in the middle of EM-1. The model predicted the variability of CO2 emissions reasonably well compared to the observations (root mean square error: 0.4-1.3gCm-2day-1, revised Willmott index: 0.16-0.55). In particular, we demonstrated that the annual reservoir surface effluxes were initially high, steeply declined in the first three years, and then steadily decreased to ~115gCm-2yr-1 with increasing reservoir age over the estimated "engineering" reservoir lifetime (i.e., 100years). Sensitivity analyses revealed that increasing air temperature stimulated CO2 emissions by enhancing CO2 production in the water column and sediment, and extending the duration of open water period over which emissions occur. Increasing the amount of terrestrial organic C flooded can enhance benthic CO2 fluxes and CO2 emissions from the reservoir water surface, but the effects were not significant over the simulation period. The model is useful for the understanding of the mechanism of C dynamics in reservoirs and could be used to assist the hydro-power industry and others interested in the role of boreal hydroelectric reservoirs as sources of greenhouse gas emissions.

Greenhouse Gas Emissions from Freshwater Reservoirs: What Does the Atmosphere See?

Freshwater reservoirs are a known source of greenhouse gas (GHG) to the atmosphere, but their quantitative significance is still only loosely con- strained. Although part of this uncertainty can be attributed to the difficulties in measuring highly variable fluxes, it is also the result of a lack of a clear accounting methodology, particularly about what constitutes new emissions and potential new sinks. In this paper, we review the main processes involved in the generation of GHG in reservoir systems and propose a simple approach to quantify the reservoir GHG footprint in terms of the net changes in GHG fluxes to the atmosphere induced by damming, that is, 'what the atmosphere sees.' The approach takes into account the pre-impoundment GHG balance of the landscape, the temporal evolution of reservoir GHG emission profile as well as the natural emissions that are displaced to or away from the reservoir site resulting from hydrological and other changes. It also clarifies the portion of the reservoir carbon burial that can potentially be considered an offset to GHG emissions.

Effects of experimental nitrogen fertilization on planktonic metabolism and CO2 flux in a hypereutrophic hardwater lake.

Hardwater lakes are common in human-dominated regions of the world and often experience pollution due to agricultural and urban effluent inputs of inorganic and organic nitrogen (N). Although these lakes are landscape hotspots for CO2 exchange and food web carbon (C) cycling, the effect of N enrichment on hardwater lake food web functioning and C cycling patterns remains unclear. Specifically, it is unknown if different eutrophication scenarios (e.g., modest non point vs. extreme point sources) yield consistent effects on auto- and heterotrophic C cycling, or how biotic responses interact with the inorganic C system to shape responses of air-water CO2 exchange. To address this uncertainty, we induced large metabolic gradients in the plankton community of a hypereutrophic hardwater Canadian prairie lake by adding N as urea (the most widely applied agricultural fertilizer) at loading rates of 0, 1, 3, 8 or 18 mg N L-1 week-1 to 3240-L, in-situ mesocosms. Over three separate 21-day experiments, all treatments of N dramatically increased phytoplankton biomass and gross primary production (GPP) two- to six-fold, but the effects of N on autotrophs plateaued at ~3 mg N L-1. Conversely, heterotrophic metabolism increased linearly with N fertilization over the full treatment range. In nearly all cases, N enhanced net planktonic uptake of dissolved inorganic carbon (DIC), and increased the rate of CO2 influx, while planktonic heterotrophy and CO2 production only occurred in the highest N treatments late in each experiment, and even in these cases, enclosures continued to in-gas CO2. Chemical effects on CO2 through calcite precipitation were also observed, but similarly did not change the direction of net CO2 flux. Taken together, these results demonstrate that atmospheric exchange of CO2 in eutrophic hardwater lakes remains sensitive to increasing N loading and eutrophication, and that even modest levels of N pollution are capable of enhancing autotrophy and CO2 in-gassing in P-rich lake ecosystems.

Surface water CO2 concentration influences phytoplankton production but not community composition across boreal lakes.

Recent experimental evidence suggests that changes in the partial pressure of CO2 (pCO2 ), in concert with nutrient fertilisation, may result in increased primary production and shifted phytoplankton community composition that favours species lacking adaptations to low CO2 environments. It is not clear whether these results apply in ambient freshwaters, which are already often supersaturated in CO2 , and where phytoplankton structure and activity are under complex control of diverse local and regional factors. Here, we use a large-scale comparative study of 69 boreal lakes to explore the influence of existing CO2 gradients (c. 50-2300 µatm) on phytoplankton community composition and biomass production. While community composition did not respond to pCO2 gradients, gross primary production was enhanced, but only in lakes already supersaturated in CO2 , demonstrating that environmental context is key in determining pCO2 -phytoplankton interactions. We further argue that increased atmospheric CO2 is unlikely to influence phytoplanktonic composition and production in northern lakes.