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Winter is a relatively under-studied season in freshwater ecology. The paucity of wintertime surveys has led to a lack of knowledge regarding microbial community activity during the winter in Lake Erie, a North American Great Lake. Viruses shape microbial communities and regulate biogeochemical cycles by acting as top-down controls, yet very few efforts have been made to examine active virus populations during the winter in Lake Erie. Furthermore, climate change-driven declines in seasonal ice cover have been shown to influence microbial community structure, but no studies have compared viral community activity between different ice cover conditions. We surveyed surface water metatranscriptomes for viral hallmark genes as a proxy for active virus populations and compared activity metrics between ice-covered and ice-free conditions from two sampled winters. Transcriptionally active viral communities were detected in both winters, spanning diverse phylogenetic clades of putative bacteriophage ( Caudoviricetes ), giant viruses ( Nucleocytoviricota , or NCLDV), and RNA viruses ( Orthornavirae ). However, viral community activity metrics revealed pronounced differences between the ice-covered and ice-free winters. Viral community composition was distinct between winters and viral hallmark gene richness was reduced in the ice-covered relative to the ice-free conditions. In addition, the observed differences in viral communities correlated with microbial community activity metrics. Overall, these findings contribute to our understanding of the viral populations that are active during the winter in Lake Erie and suggest that viral community activity may be associated with ice cover extent.IMPORTANCEAs seasonal ice cover is projected to become increasingly rare on large temperate lakes, there is a need to understand how microbial communities might respond to changing ice conditions. Although it is widely recognized that viruses impact microbial community structure and function, there is little known regarding wintertime viral activity or the relationship between viral activity and ice cover extent. Our metatranscriptomic analyses indicated that viruses were transcriptionally active in the winter surface waters of Lake Erie. These findings also expanded the known diversity of viral lineages in the Great Lakes. Notably, viral community activity metrics were significantly different between the two sampled winters. The pronounced differences we observed in active viral communities between the ice-covered and ice-free samples merit further research regarding how viral communities will function in future, potentially ice-free, freshwater systems., Competing Interests: The authors declare no conflict of interest.

How climate change will affect the community dynamics and functionality of lake ecosystems during winter is still little understood. This is also true for phytoplankton in seasonally ice-covered temperate lakes which are particularly vulnerable to the presence or absence of ice. We examined changes in pelagic phytoplankton winter community structure in a north temperate lake (Müggelsee, Germany), covering 18 winters between 1995 and 2013. We tested how phytoplankton taxa composition varied along a winter-severity gradient and to what extent winter severity shaped the functional trait composition of overwintering phytoplankton communities using multivariate statistical analyses and a functional trait-based approach. We hypothesized that overwintering phytoplankton communities are dominated by taxa with trait combinations corresponding to the prevailing winter water column conditions, using ice thickness measurements as a winter-severity indicator. Winter severity had little effect on univariate diversity indicators (taxon richness and evenness), but a strong relationship was found between the phytoplankton community structure and winter severity when taxon trait identity was taken into account. Species responses to winter severity were mediated by the key functional traits: motility, nutritional mode, and the ability to form resting stages. Accordingly, one or the other of two functional groups dominated the phytoplankton biomass during mild winters (i.e., thin or no ice cover; phototrophic taxa) or severe winters (i.e., thick ice cover; exclusively motile taxa). Based on predicted milder winters for temperate regions and a reduction in ice-cover durations, phytoplankton communities during winter can be expected to comprise taxa that have a relative advantage when the water column is well mixed (i.e., need not be motile) and light is less limiting (i.e., need not be mixotrophic). A potential implication of this result is that winter severity promotes different communities at the vernal equinox, which may have different nutritional quality for the next trophic level and ecosystem-scale effects., (© 2015 John Wiley & Sons Ltd.)

Carbon and nutrient cycles in large temperate lakes such as Lake Erie are primarily driven by phototrophic and heterotrophic microorganisms, although our understanding of these is often constrained to late spring through summer due to logistical constraints. During periods of > 90% ice cover in February of 2008, 2009, and 2010, we collected samples from an icebreaker for an examination of bacterial production as well as microbial community structure. In comparison with summer months (August 2002 and 2010), we tested hypotheses concerning seasonal changes in microbial community diversity and production. Bacterial production estimates were c. 2 orders of magnitude higher (volume normalized) in summer relative to winter. Our observations further demonstrate that the microbial community, including single-celled phototrophs, varied in composition between August and February. Sediment traps deployed and collected over a 3 year period (2008-2011) confirmed that carbon export was ongoing and not limiting winter production. The results support the notion that active primary producers in winter months export carbon to the sediments that is not consumed until the warmer seasons. The establishment of this linkage is a critical observation in efforts to understand the extent and severity of annual summertime formations of a zone of regional hypoxia in Lake Erie., (© 2013 Federation of European Microbiological Societies. Published by John Wiley & Sons Ltd. All rights reserved.)

Global surface freshwater primarily resides in lakes, with the overwhelming majority found in Earth’s largest lakes, thus understanding potential climate change effects in these large lakes is critical. In dimictic lakes, climate change has extended the duration of summer thermal stratification and reduced the length of the ice season. These changes are relatively straightforward to evaluate in smaller, inland lakes. However, in large lakes, such as the North American Great Lakes, temporally intermittent and spatially heterogeneous ice cover, and spatial thermal heterogeneity limit the utility of simple ice on–off or mixing classifications; therefore, assessing how climate change is impacting winter conditions in large lakes is challenging. Here, we use in-situ and satellite-derived surface water temperature observations from the North American Great Lakes to overcome these limitations and show that warming air temperatures are driving reductions in the number of winter days, collectively those with either ice cover or inverse thermal stratification, in favor of increases in isothermal conditions for the period 1995–2023. We find that on average the Great Lakes are experiencing a loss of 14 winter days per decade. Our results demonstrate how climate change has yielded disproportionate changes in the annual thermal cycle and mixing conditions of Earth’s largest freshwater system and signals the potential for fundamental ecosystem shifts due to a loss of winter.

The frequency and duration of lake ice cover is rapidly declining in the Northern Hemisphere. Limited research in oligotrophic and mesotrophic lakes suggests that extended periods of ice cover influence nitrogen (N) cycling by promoting nitrate (NO3−) accumulation. However, ice cover impacts on N cycling in shallow, high-nutrient, eutrophic lakes remains poorly understood. To understand drivers of under-ice water column N concentrations, we examined concurrent under-ice N concentration, hydro-meterological, and physicochemical time series from two shallow eutrophic systems during sustained cold and thaw periods. We compared data from both lakes during a historically cold winter to assess how different lake-watershed physical configurations and algal biomass affected under-ice N cycling. We also used time series data from consecutive winters to assess the influence of a mild versus a historically cold winter on under-ice N cycling in one of the lakes. We found that ice cover promoted NO3− depletion when sustained cold disconnected lakes from watershed loading, but the amount of depletion varied between lakes and was enhanced during the colder winter. Thaw events increased NO3− concentrations compared to late winter and altered the concentration and distribution of N species in the water column, but the degree and nature of increased NO3− varied with thaw severity, the source of meltwater, timing, and lake-watershed morphometry. Our results suggest that winters with shorter ice duration and more thaw events may result in less NO3− depletion and higher peak NO3− concentrations in shallow eutrophic lakes, with potential implications for N cycling and phytoplankton ecology. [ABSTRACT FROM AUTHOR]

The Qinghai‐Tibet Plateau possesses the largest alpine lake system, which plays a crucial role in the land‐atmosphere interaction. We report first observations on the thermal and radiation regime under ice of the largest freshwater lake of the Plateau. The results reveal that freshwater lakes on the Tibetan Plateau fully mix under ice. Due to strong solar heating, water temperatures increase above the maximum density value 1–2 months before the ice break, forming stable thermal stratification with subsurface temperatures >6°C. The resulting heat flow from water to ice makes a crucial contribution to ice cover melt. After the ice breakup, the accumulated heat is released into the atmosphere during 1–2 days, increasing lake‐atmosphere heat fluxes up to 500 W m−2. The direct biogeochemical consequences of the deep convective mixing are aeration of the deep lake waters and upward supply of nutrients to the upper photic layer. Plain Language Summary: The Qinghai‐Tibet Plateau possesses the largest alpine lake system, which plays a crucial role in the land‐atmosphere interaction. Data on thermal properties of Tibetan lakes during the ice‐covered season are extremely scarce. The first observations on the thermal and radiation regime under ice of the largest freshwater lake of the Plateau reveal that freshwater (and apparently the majority of brackish) lakes gain an extremely large amount of solar radiation penetrating the highly transparent ice cover. As a result, lakes fully mix under ice and get heated up to >6°C. The accumulated heat makes a crucial contribution to ice cover melt. The quick release of the heat to the atmosphere during 1–2 days after the ice breakup strongly increases lake‐atmosphere heat fluxes, potentially affecting regional weather conditions. Warm and well‐mixed conditions are favorable for aquatic life in the extreme environment of Tibet. Key Points: An abnormal thermal regime under the ice cover of Tibetan lakes is revealedThe lakes get heated above their maximum density temperature by extremely high level of solar radiation penetrating the ice coverThe stored heat shortens the ice‐covered period and is quickly released into the atmosphere after ice‐off, affecting local climate [ABSTRACT FROM AUTHOR]

Dissolved oxygen controls important processes in lakes, from chemical reactions to organism community structure and metabolism. In shallow lakes, small volumes allow for large fluctuations in dissolved oxygen concentrations, and the oxygen regime can greatly affect ecosystem‐scale processes. We used high frequency dissolved oxygen measurements to examine differences in oxygen regimes between two alternative stable states that occur in shallow lakes. We compared annual oxygen regimes in four macrophyte‐dominated, clear state lakes to four phytoplankton‐dominated, turbid state lakes by quantifying oxygen concentrations, anoxia frequency, and measures of whole‐lake metabolism. Oxygen regimes were not significantly different between lake states throughout the year except for during the winter under‐ice period. During winter, clear lakes had less oxygen, higher frequency of anoxic periods, and higher oxygen depletion rates. Winter oxygen depletion rates correlated positively with peak summer macrophyte biomass. Due to lower levels of oxygen, clear shallow lakes may experience anoxia more often and for longer duration during the winter, increasing the likelihood of fish winterkills. These observations have important implications for shallow lake management, which typically focuses efforts on maintaining the clearwater state. Plain Language Summary: In lakes, the amount of oxygen dissolved in the water has a profound impact on lake processes, from chemical reactions to the varieties and quantities of organisms present. In shallow lakes, the amount of dissolved oxygen can vary greatly due to the differences in the rates of production, mostly through photosynthesis, and consumption, mostly through respiration. In this study, we compared dissolved oxygen availability seasonally between two common states found in shallow lakes; a turbid, low clarity state dominated by phytoplankton, and a clear state dominated by submersed aquatic plants. Patterns of oxygen were similar between the two lakes states in all seasons except winter. During the winter under‐ice period, clear lakes had significantly less oxygen compared to turbid lakes, and lost oxygen at a faster rate through the winter. The lower levels of oxygen in clear lakes during the winter could affect many lake processes, such as the winterkill of fish. Lake managers typically try to maintain shallow lakes in the clear state because of better water quality and increased wildlife diversity. Winter fish kills could help maintain lakes in the clear state, but may also select for rough fish that can drive shifts to the turbid state. Key Points: Winter oxygen regimes differed between clear and turbid shallow lakesClear shallow lakes had significantly higher oxygen depletion rates under ice cover than turbid shallow lakesOxygen depletion rates were highly correlated with summer macrophyte biomass [ABSTRACT FROM AUTHOR]

Lakes are traditionally classified based on their thermal regime and trophic status. While this classification adequately captures many lakes, it is not sufficient to understand seasonally ice‐covered lakes, the most common lake type on Earth. We describe the inverse thermal stratification in 19 highly varying lakes and derive a model that predicts the temperature profile as a function of wind stress, area, and depth. The results suggest an additional subdivision of seasonally ice‐covered lakes to differentiate underice stratification. When ice forms in smaller and deeper lakes, inverse stratification will form with a thin buoyant layer of cold water (near 0°C) below the ice, which remains above a deeper 4°C layer. In contrast, the entire water column can cool to ∼0°C in larger and shallower lakes. We suggest these alternative conditions for dimictic lakes be termed "cryostratified" and "cryomictic." Plain Language Summary: Most mid and high latitude lakes are seasonally ice‐covered and have only been classified based on the thermal structure and trophic status during the open‐water season in summer. However, limited temperatures observations in these ice‐covered lakes suggest that there is a wide range of thermal structures over winter. We developed an analytical model to predict the average water temperature at the time of ice formation based on the strength of the surface winds, the area of the lake, and the maximum depth of the lake. Using both the analytical model and water temperature data from 19 different lakes in North America, Europe, and Asia, we found that the time of ice formation in lakes that are large or experience strong winds were later compared to lakes that are small or experience weak winds. The larger and windier lakes are also colder (0°C∼2°C) than smaller and calmer lakes (2°C∼4°C) at the time of ice formation. This suggests that these seasonally ice‐covered lakes can be subdivided into two additional classes during winter. The analytical model and the new categorization have important consequences for understanding fish habitat under the ice and the potential effects of climate change on these seasonally ice‐covered lakes. Key Points: Standard classifications of dimictic lakes do not consider how variable the initial thermal stratification can be under winter lake iceLakes that are shallow or windy can cool to near 0°C–1°C before ice forms and are weakly stratified, which we term "cryomictic"Deeper lakes or those with calmer winds, result in ice forming just above deeper waters of 3°C–4°C, which we term "cryostratified" [ABSTRACT FROM AUTHOR]

Winter is a long period of the annual cycle of many lakes in the northern hemisphere. Low irradiance, ice, and snow cover cause poor light penetration into the water column of these lakes. Therefore, in northern lakes, respiration often exceeds primary production leading to low dissolved oxygen concentrations. This study aimed to quantify under‐ice metabolic processes during winter in an arid zone lake with little snow cover.This study was carried out in a mid‐latitude lake in Inner Mongolia, northern China. The study lake receives relatively high incoming solar radiation on the ice in mid‐winter, and radiation can penetrate down to the bottom sediment as the lake is shallow and the ice lacks snow cover.Primary production and respiration were estimated during two winters using high‐frequency sensor measurements of dissolved oxygen. To quantify under‐ice metabolic processes, sensors were deployed to different depths. During both winters, sensors collected data every 10 min over several weeks.The amount of solar radiation controlled photosynthesis under ice; temperature and photosynthesis together appeared to control respiration. The balance between gross primary production and ecosystem respiration was especially sensitive to changes in snow cover, and the balance between P and R decreased.Our data suggest that photosynthesis by plankton, submerged plants, and epiphytic algae may continue over winter in shallow lakes in mid‐latitudes when there is no snow cover on the ice, as may occur in arid climates. The continuation of photosynthesis under ice buffers against dissolved oxygen depletion and prevents consequent harmful ecosystem effects. [ABSTRACT FROM AUTHOR]

[Display omitted] • Climate change results in a decrease of the ice cover and duration in Northern Hemisphere lakes. • Warming resulted in ice decline and will continue to negatively affect lake ice in the future. • Strong autumnal winds can accelerate ice formation but this effect apparently depends on the mixing. • Rainfalls either enhance freezing in fall or accelerate ice-loss in spring. • Under continuous emissions lakes will further lose their ice and experience ice-free winters more frequently. Northern Hemisphere lakes are losing their ice cover due to climate change. Here we explored six decades of observational data (1961–2017) showing trends in air temperature, wind speed and precipitation over northern Poland, as well as changes in the ice conditions for five lakes with different morphometry. We evaluated whether and to what extent climatic effects, including atmospheric warming, changing wind speed and rainfall during fall and winter, influence ice conditions in morphometrically different lakes in Northern Poland. Our analysis demonstrated that ice cover duration and thickness decreased at rates of 5.4 days decade−1 and 2.5 cm decade−1, respectively. Ice conditions were influenced (65–75%) by the direct effects of air temperature change and to some extent by an interaction of warming with wind speed and precipitation (5–10%). While stronger autumnal winds result in longer ice cover duration, the effect of precipitation is bimodal with either an enhancement of ice formation by autumnal rain or accelerated ice loss during spring. To project future changes in ice conditions, we used a 1D hydrodynamic lake model forced with four climate model projections under low, medium and high Representative Concentration Pathway (RCP) scenarios. Our simulations demonstrate that current ice conditions will stabilize under the low emission scenario (RCP 2.6) but decrease under both the medium and high emission scenarios (RCP 6.0 and 8.5). During the 21st century, the lakes are projected to lose their ice at a rate between 4.5- and 10-days decade−1 and ice thickness will decrease by between 3.0 and 5.0 cm decade−1. The rate of change will be greater in smaller rather than larger lakes and more so for those situated further inland. The probability of ice-free winters will increase for all lakes and among all future scenarios by between 4 and 69% with the highest potential frequency of ice-free winters in smaller and deeper but relatively wind-exposed lakes. [ABSTRACT FROM AUTHOR]