Millions of lakes worldwide are distributed at latitudes or elevations resulting in the formation of lake ice during winter. Lake ice affects the transfer of energy, heat, light, and material between lakes and their surroundings creating an environment dramatically different from open-water conditions. While this fundamental restructuring leads to distinct gradients in ions, dissolved gases, and nutrients throughout the water column, surprisingly little is known about the resulting effects on ecosystem processes and food webs, highlighting the lack of a general limnological framework that characterizes the structure and function of lakes under a gradient of ice cover. Drawing from the literature and three novel case studies, we present the Lake Ice Continuum Concept (LICC) as a model for understanding how key aspects of the physical, chemical, and ecological structure and function of lakes vary along a continuum of winter climate conditions mediated by ice and snow cover. We examine key differences in energy, redox, and ecological community structure and describe how they vary in response to shifts in physical mixing dynamics and light availability for lakes with ice and snow cover, lakes with clear ice alone, and lakes lacking winter ice altogether. Global change is driving ice covered lakes toward not only warmer annual average temperatures but also reduced, intermittent or no ice cover. The LICC highlights the wide range of responses of lakes to ongoing climate-driven changes in ice cover and serves as a reminder of the need to understand the role of winter in the annual aquatic cycle.Plain Language Summary Millions of lakes worldwide freeze during winter. The formation of lake ice dramatically alters lakes by isolating them from their surrounding landscape and atmosphere. The thickness and optical qualities of ice and snow regulate the amount of solar radiation entering lakes, while shielding them from wind energy. Consequently, lake ice is an important factor regulating physical mixing dynamics within lakes, and structuring vertical thermal and chemical gradients. Although organisms from bacteria to fish have adapted to the winter environment, we lack a comprehensive understanding of how variation in lake ice cover conditions affects fundamental ecosystem processes or food web structure. Here, we combine a synthesis of the current literature with three novel case studies to develop the Lake Ice Continuum Concept as a framework for understanding how key aspects of the physical, chemical, and ecological structure and function of lakes vary along a continuum of energy inputs mediated by winter climate. This framework is useful for understanding changes associated with seasonal ice dynamics and for predicting how lakes may respond to climate change.

Salinization of freshwater is increasingly observed in regions where chloride de-icing salts are applied to the roads in winter, but little is known about the effects on microbial communities. In this study, we analyzed the planktonic microbiomes of four lakes that differed in degree of urbanization, eutrophication and salinization, from an oligotrophic reference lake with no surrounding roads, to a eutrophic, salinized lake receiving runoff from a highway. We tested the hypothesis that an influence of road salts would be superimposed on the effects of season and trophic status. We evaluated the microbial community structure by 16S rRNA sequencing for Bacteria, and by four methods for eukaryotes: 16S rRNA chloroplast analysis, 18S rRNA sequencing, photosynthetic pigment analysis and microscopy. Consistent with our hypothesis, chloride and total nitrogen concentrations were among the most important statistical factors explaining the differences in taxonomic composition. These factors were positively correlated with the abundance of cryptophytes, haptophytes, and cyanobacteria. Ice-cover was also a major structuring factor, with clear differences between the winter communities and those of the open-water period. Nitrifying and methane oxidizing bacteria were more abundant in winter, suggesting the importance of anaerobic sediment processes and release of reduced compounds into the ice-covered water columns. The four methods for eukaryotic analysis provided complementary information. The 18S rRNA observations were strongly influenced by the presence of ribosome-rich ciliates, but revealed a much higher degree of taxonomic richness and greater separation of lakes, seasonal changes and potential salinity effects than the other methods.

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