Freshwater lakes and reservoirs play a disproportionate role in the global organic carbon (OC) budget, as active sites for carbon processing and burial. Associations between OC and iron (Fe) are hypothesized to contribute substantially to the stabilization of OC in sediment, but the magnitude of freshwater Fe-OC complexation remains unresolved. Moreover, global declines in bottom-water oxygen concentrations have the potential to alter OC and Fe cycles in multiple ways, and the net effects of low-oxygen (hypoxic) conditions on OC and Fe are poorly characterized. Here, we measured the pool of Fe-bound OC (Fe-OC) in surficial sediments from two eutrophic reservoirs, and we paired whole-ecosystem experiments with sediment incubations to determine the effects of hypoxia on OC and Fe cycling over multiple timescales. Our experiments demonstrated that short (2–4 week) periods of hypoxia can increase aqueous Fe and OC concentrations while decreasing OC and Fe-OC in surficial sediment by 30%. However, exposure to seasonal hypoxia over multiple years was associated with a 57% increase in sediment OC and no change in sediment Fe-OC. These results suggest that the large sediment Fe-OC pool (~30% of sediment OC in both reservoirs) contains both oxygen-sensitive and oxygen-insensitive fractions, and over multiannual timescales OC respiration rates may play a more important role in in determining the effect of hypoxia on sediment OC than Fe-OC dissociation. Consequently, we anticipate that global declines in oxygen concentrations will alter OC and Fe cycling, with the direction and magnitude of effects dependent upon the duration of hypoxia.

Temperate reservoirs and lakes worldwide are experiencing decreases in ice cover, which will likely alter the net balance of gross primary production (GPP) and respiration (R) in these ecosystems. However, most metabolism studies to date have focused on summer dynamics, thereby excluding winter dynamics from annual metabolism budgets. To address this gap, we analyzed six years of year-round high-frequency dissolved oxygen data to estimate daily rates of net ecosystem production (NEP), GPP, and R in a eutrophic, dimictic reservoir that has intermittent ice cover. Over six years, the reservoir exhibited slight heterotrophy during both summer and winter. We found winter and summer metabolism rates to be similar: summer NEP had a median rate of -0.06 mg O2 L-1 day-1 (range: -15.86 to 3.20 mg O2 L-1 day-1), while median winter NEP was -0.02 mg O2 L-1 day-1 (range: -8.19 to 0.53 mg O2 L-1 day-1). Despite large differences in the duration of ice cover among years, there were minimal differences in NEP among winters. Overall, the inclusion of winter data had a limited effect on annual metabolism estimates, likely due to short winter periods in this reservoir (ice durations 0–35 days) relative to higher-latitude lakes. Our work reveals a smaller difference between winter and summer NEP than in lakes with continuous ice cover. Ultimately, our work underscores the importance of studying full-year metabolism dynamics in a range of aquatic ecosystems to help anticipate the effects of declining ice cover across lakes worldwide.

Water level drawdowns are increasingly common in lakes and reservoirs worldwide as a result of both climate change and water management. Drawdowns can have direct effects on physical properties of a waterbody (e.g., by altering stratification and light dynamics), which can interact to modify the waterbody’s biology and chemistry. However, the ecosystem-level effects of drawdown remain poorly characterized in small, thermally-stratified reservoirs, which are common in many regions of the world. Here, we intensively monitored a small eutrophic reservoir for two years, including before, during, and after a month-long drawdown that reduced total reservoir volume by 36%. During drawdown, stratification strength (maximum buoyancy frequency) and surface phosphate concentrations both increased, contributing to a substantial surface phytoplankton bloom. The peak in phytoplankton biomass was followed by cascading changes in surface water chemistry associated with bloom degradation, with sequential peaks in dissolved organic carbon, dissolved carbon dioxide, and ammonium concentrations that were up to an order of magnitude higher than the previous year. Dissolved oxygen concentrations substantially decreased in the surface waters during drawdown (to 41% saturation), which was associated with increased total iron and manganese concentrations. Combined, our results illustrate how changes in water level can have cascading effects on coupled physical, chemical, and biological processes. As climate change and water management continue to increase the frequency of drawdowns in lakes worldwide, our results highlight the importance of characterizing how water level variability can alter complex in-lake ecosystem processes, thereby affecting water quality.

Declining oxygen concentrations in the deep waters of lakes worldwide pose a pressing environmental and societal challenge. Existing theory suggests that low deep-water dissolved oxygen (DO) concentrations could trigger a positive feedback through which anoxia (i.e., very low DO) during a given summer begets increasingly severe occurrences of anoxia in following summers. Specifically, anoxic conditions can promote nutrient release from sediments, thereby stimulating phytoplankton growth, and subsequent phytoplankton decomposition can fuel heterotrophic respiration, resulting in increased spatial extent and duration of anoxia. However, while the individual relationships in this feedback are well established, to our knowledge there has not been a systematic analysis within or across lakes that simultaneously demonstrates all of the mechanisms necessary to produce a positive feedback that reinforces anoxia. Here, we compiled data from 656 widespread temperate lakes and reservoirs to analyze the proposed Anoxia Begets Anoxia (ABA) feedback. Lakes in the dataset span a broad range of surface area (1–126,909 ha), maximum depth (6–370 m), and morphometry, with a median time series duration of 30 years at each lake. Using linear mixed models, we found support for each of the positive feedback relationships between anoxia, phosphorus concentrations, chlorophyll-a concentrations, and oxygen demand across the 656-lake dataset. Likewise, we found further support for these relationships by analyzing time series data from individual lakes. Our results indicate that the strength of these feedback relationships may vary with lake-specific characteristics: for example, we found that surface phosphorus concentrations were more positively associated with chlorophyll-a in high-phosphorus lakes, and oxygen demand had a stronger influence on the extent of anoxia in deep lakes. Taken together, these results support the existence of a positive feedback that could magnify the effects of climate change and other anthropogenic pressures driving the development of anoxia in lakes around the world.

Scientific Significance StatementSummer bottom-water temperature and dissolved oxygen dynamics play a critical role in shaping water quality in many temperate lakes. Counterintuitively, it is possible that spring air temperatures exert a stronger influence on summer bottom-water dynamics than summer air temperatures—a phenomenon we refer to as “seasonal ecological memory.” In our analysis of 615 temperate lakes, we found support for a seasonal ecological memory effect that helps explain why trends in summer bottom-water temperature diverge from trends in summer air temperature, thereby improving our ability to predict the effects of climate change on lake water quality dynamics.

Oxygen availability is decreasing in many lakes and reservoirs worldwide, raising the urgency for understanding how anoxia (low oxygen) affects coupled biogeochemical cycling, which has major implications for water quality, food webs, and ecosystem functioning. Although the increasing magnitude and prevalence of anoxia has been documented in freshwaters globally, the challenges of disentangling oxygen and temperature responses have hindered assessment of the effects of anoxia on carbon, nitrogen, and phosphorus concentrations, stoichiometry (chemical ratios), and retention in freshwaters. The consequences of anoxia are likely severe and may be irreversible, necessitating ecosystem-scale experimental investigation of decreasing freshwater oxygen availability. To address this gap, we devised and conducted REDOX (the Reservoir Ecosystem Dynamic Oxygenation eXperiment), an unprecedented, seven-year experiment in which we manipulated and modeled bottom-water (hypolimnetic) oxygen availability at the whole-ecosystem scale in a eutrophic reservoir. Seven years of data reveal that anoxia significantly increased hypolimnetic carbon, nitrogen, and phosphorus concentrations and altered elemental stoichiometry by factors of 2-5 relative to oxic periods. Importantly, prolonged summer anoxia increased nitrogen export from the reservoir by six-fold and changed the reservoir from a net sink to a net source of phosphorus and organic carbon downstream. While low oxygen in freshwaters is thought of as a response to land use and climate change, results from REDOX demonstrate that low oxygen can also be a driver of major changes to freshwater biogeochemical cycling, which may serve as an intensifying feedback that increases anoxia in downstream waterbodies. Consequently, as climate and land use change continue to increase the prevalence of anoxia in lakes and reservoirs globally, it is likely that anoxia will have major effects on freshwater carbon, nitrogen, and phosphorus budgets as well as water quality and ecosystem functioning.

1. Freshwater phytoplankton communities are currently experiencing multiple global change stressors, including increasing frequency and intensity of storms. An important mechanism by which storms affect lake and reservoir phytoplankton is by altering the water column’s thermal structure (e.g., changes to thermocline depth). However, little is known about the effects of intermittent thermocline deepening on phytoplankton community vertical distribution and composition or the consistency of phytoplankton responses to varying frequency of these disturbances over multiple years. 2. We conducted whole-ecosystem thermocline deepening manipulations in a small reservoir. We used an epilimnetic mixing system to experimentally deepen the thermocline in two summers, simulating potential responses to storms, and did not manipulate thermocline depth in two succeeding summers. We collected weekly depth profiles of water temperature, light, nutrients, and phytoplankton biomass as well as discrete samples to assess phytoplankton community composition. We then used time-series analysis and multivariate ordination to assess the effects of intermittent thermocline deepening due to both our experimental manipulations and naturally-occurring storms on phytoplankton community structure. 3. We observed inter-annual and intra-annual variability in phytoplankton community response to thermocline deepening. We found that peak phytoplankton biomass was significantly deeper in years with a higher frequency of thermocline deepening events (i.e., years with both manipulations and natural storms) due to weaker thermal stratification and deeper depth distributions of soluble reactive phosphorus. Furthermore, we found that the depth of peak phytoplankton biomass was linked to phytoplankton community composition, with certain taxa being associated with deep or shallow biomass peaks, often according to functional traits such as optimal growth temperature, mixotrophy, and low-light tolerance. 4. Our results demonstrate that abrupt thermocline deepening due to water column mixing affects both phytoplankton depth distribution and community structure via alteration of physical and chemical gradients. In addition, our work supports previous research that phytoplankton depth distribution and community composition interact at inter-annual and intra-annual timescales. 5. Variability in the inter-annual and intra-annual responses of phytoplankton to abrupt thermocline deepening indicates that antecedent conditions and the seasonal timing of surface water mixing may mediate these responses. Our findings emphasize that phytoplankton depth distributions are sensitive to global change stressors and effects on depth distributions should be taken into account when predicting phytoplankton responses to increased storms under global change.

Globally-significant quantities of carbon (C), nitrogen (N), and phosphorus (P) enter freshwater reservoirs each year. These inputs can be buried in sediments, respired, taken up by organisms, emitted to the atmosphere, or exported downstream. While much is known about reservoir-scale biogeochemical processing, less is known about spatial and temporal variability of biogeochemistry within a reservoir along the continuum from inflowing streams to the dam. To address this gap, we examined longitudinal variability in surface water biogeochemistry (C, N, and P) in two small reservoirs throughout a thermally-stratified season. We sampled total and dissolved fractions of C, N, and P, and chlorophyll-a from each reservoir’s major inflows to the dam. We found that time was generally a more important driver of heterogeneity in biogeochemical concentrations than space. However, dissolved nutrient and organic carbon concentrations had high site-to-site variability within both reservoirs, potentially as a result of shifting biological activity or environmental conditions. When considering spatially explicit processing, we found that certain locations within the reservoir, most often the stream-reservoir interface, acted as ‘hotspots’ of change in biogeochemical concentrations. Our study suggests that spatially explicit metrics of biogeochemical processing could help constrain the role of reservoirs in C, N, and P cycles in the landscape. Ultimately, our results highlight that biogeochemical heterogeneity in small reservoirs is driven more by seasonality than longitudinal spatial gradients, and that some sites within reservoirs play critically important roles in whole-ecosystem biogeochemical processing.