Widespread deoxygenation of temperate lakes.

The concentration of dissolved oxygen in aquatic systems helps to regulate biodiversity1,2, nutrient biogeochemistry3, greenhouse gas emissions4, and the quality of drinking water5. The long-term declines in dissolved oxygen concentrations in coastal and ocean waters have been linked to climate warming and human activity6,7, but little is known about the changes in dissolved oxygen concentrations in lakes. Although the solubility of dissolved oxygen decreases with increasing water temperatures, long-term lake trajectories are difficult to predict. Oxygen losses in warming lakes may be amplified by enhanced decomposition and stronger thermal stratification8,9 or oxygen may increase as a result of enhanced primary production10. Here we analyse a combined total of 45,148 dissolved oxygen and temperature profiles and calculate trends for 393 temperate lakes that span 1941 to 2017. We find that a decline in dissolved oxygen is widespread in surface and deep-water habitats. The decline in surface waters is primarily associated with reduced solubility under warmer water temperatures, although dissolved oxygen in surface waters increased in a subset of highly productive warming lakes, probably owing to increasing production of phytoplankton. By contrast, the decline in deep waters is associated with stronger thermal stratification and loss of water clarity, but not with changes in gas solubility. Our results suggest that climate change and declining water clarity have altered the physical and chemical environment of lakes. Declines in dissolved oxygen in freshwater are 2.75 to 9.3 times greater than observed in the world's oceans6,7 and could threaten essential lake ecosystem services2,3,5,11.

Disentangling the role of light and nutrient limitation on bacterivory by mixotrophic nanoflagellates.

Many phytoplankton taxa function on multiple trophic levels by combining photosynthesis and ingestion of bacteria, termed mixotrophy. Despite the recognition of mixotrophy as a universal functional trait, we have yet to fully resolve how environmental conditions influence community grazing rates in situ. A microcosm study was used to assess bacterivory by mixotrophic nanoflagellates following nutrient enrichment and light attenuation in a temperate lake. We found contrasting results based on assessment of mixotroph abundance or bacterivory. Despite an interactive effect of nutrient enrichment and light attenuation on mixotroph abundance, significant differences within light treatments were observed only after enrichment with P or N + P. The greatest abundance of mixotrophs across treatments occurred under co-nutrient enrichment with full exposure to irradiance. However, bacterivory by mixotrophic nanoflagellates was greatest under shaded conditions after either N or P enrichment. We suggest that PAR availability dampened the stimulatory effect of nutrient limitation, and bacterivory supplemented a suboptimal photosynthetic environment. In a saturating light regime, the mixotrophic community was less driven to ingest bacteria because photosynthesis was able to satisfy energetic demands. These findings quantify community bacterivory in response to environmental drivers that may characterize future ecosystem conditions and highlight the importance of considering grazing rates in conjunction with abundance of mixotrophic protists.

The success of the Montreal Protocol in mitigating interactive effects of stratospheric ozone depletion and climate change on the environment.

The Montreal Protocol and its Amendments have been highly effective in protecting the stratospheric ozone layer, preventing global increases in solar ultraviolet-B radiation (UV-B; 280-315 nm) at Earth's surface, and reducing global warming. While ongoing and projected changes in UV-B radiation and climate still pose a threat to human health, food security, air and water quality, terrestrial and aquatic ecosystems, and construction materials and fabrics, the Montreal Protocol continues to play a critical role in protecting Earth's inhabitants and ecosystems by addressing many of the United Nations Sustainable Development Goals.

Insects in high-elevation streams: Life in extreme environments imperiled by climate change.

Climate change is altering conditions in high-elevation streams worldwide, with largely unknown effects on resident communities of aquatic insects. Here, we review the challenges of climate change for high-elevation aquatic insects and how they may respond, focusing on current gaps in knowledge. Understanding current effects and predicting future impacts will depend on progress in three areas. First, we need better descriptions of the multivariate physical challenges and interactions among challenges in high-elevation streams, which include low but rising temperatures, low oxygen supply and increasing oxygen demand, high and rising exposure to ultraviolet radiation, low ionic strength, and variable but shifting flow regimes. These factors are often studied in isolation even though they covary in nature and interact in space and time. Second, we need a better mechanistic understanding of how physical conditions in streams drive the performance of individual insects. Environment-performance links are mediated by physiology and behavior, which are poorly known in high-elevation taxa. Third, we need to define the scope and importance of potential responses across levels of biological organization. Short-term responses are defined by the tolerances of individuals, their capacities to perform adequately across a range of conditions, and behaviors used to exploit local, fine-scale variation in abiotic factors. Longer term responses to climate change, however, may include individual plasticity and evolution of populations. Whether high-elevation aquatic insects can mitigate climatic risks via these pathways is largely unknown.

Key rules of life and the fading cryosphere: Impacts in alpine lakes and streams.

Alpine regions are changing rapidly due to loss of snow and ice in response to ongoing climate change. While studies have documented ecological responses in alpine lakes and streams to these changes, our ability to predict such outcomes is limited. We propose that the application of fundamental rules of life can help develop necessary predictive frameworks. We focus on four key rules of life and their interactions: the temperature dependence of biotic processes from enzymes to evolution; the wavelength dependence of the effects of solar radiation on biological and ecological processes; the ramifications of the non-arbitrary elemental stoichiometry of life; and maximization of limiting resource use efficiency across scales. As the cryosphere melts and thaws, alpine lakes and streams will experience major changes in temperature regimes, absolute and relative inputs of solar radiation in ultraviolet and photosynthetically active radiation, and relative supplies of resources (e.g., carbon, nitrogen, and phosphorus), leading to nonlinear and interactive effects on particular biota, as well as on community and ecosystem properties. We propose that applying these key rules of life to cryosphere-influenced ecosystems will reduce uncertainties about the impacts of global change and help develop an integrated global view of rapidly changing alpine environments. However, doing so will require intensive interdisciplinary collaboration and international cooperation. More broadly, the alpine cryosphere is an example of a system where improving our understanding of mechanistic underpinnings of living systems might transform our ability to predict and mitigate the impacts of ongoing global change across the daunting scope of diversity in Earth's biota and environments.

The interactive effects of stratospheric ozone depletion, UV radiation, and climate change on aquatic ecosystems.

This assessment summarises the current state of knowledge on the interactive effects of ozone depletion and climate change on aquatic ecosystems, focusing on how these affect exposures to UV radiation in both inland and oceanic waters. The ways in which stratospheric ozone depletion is directly altering climate in the southern hemisphere and the consequent extensive effects on aquatic ecosystems are also addressed. The primary objective is to synthesise novel findings over the past four years in the context of the existing understanding of ecosystem response to UV radiation and the interactive effects of climate change. If it were not for the Montreal Protocol, stratospheric ozone depletion would have led to high levels of exposure to solar UV radiation with much stronger negative effects on all trophic levels in aquatic ecosystems than currently experienced in both inland and oceanic waters. This "world avoided" scenario that has curtailed ozone depletion, means that climate change and other environmental variables will play the primary role in regulating the exposure of aquatic organisms to solar UV radiation. Reductions in the thickness and duration of snow and ice cover are increasing the levels of exposure of aquatic organisms to UV radiation. Climate change was also expected to increase exposure by causing shallow mixed layers, but new data show deepening in some regions and shoaling in others. In contrast, climate-change related increases in heavy precipitation and melting of glaciers and permafrost are increasing the concentration and colour of UV-absorbing dissolved organic matter (DOM) and particulates. This is leading to the "browning" of many inland and coastal waters, with consequent loss of the valuable ecosystem service in which solar UV radiation disinfects surface waters of parasites and pathogens. Many organisms can reduce damage due to exposure to UV radiation through behavioural avoidance, photoprotection, and photoenzymatic repair, but meta-analyses continue to confirm negative effects of UV radiation across all trophic levels. Modeling studies estimating photoinhibition of primary production in parts of the Pacific Ocean have demonstrated that the UV radiation component of sunlight leads to a 20% decrease in estimates of primary productivity. Exposure to UV radiation can also lead to positive effects on some organisms by damaging less UV-tolerant predators, competitors, and pathogens. UV radiation also contributes to the formation of microplastic pollutants and interacts with artificial sunscreens and other pollutants with adverse effects on aquatic ecosystems. Exposure to UV-B radiation can decrease the toxicity of some pollutants such as methyl mercury (due to its role in demethylation) but increase the toxicity of other pollutants such as some pesticides and polycyclic aromatic hydrocarbons. Feeding on microplastics by zooplankton can lead to bioaccumulation in fish. Microplastics are found in up to 20% of fish marketed for human consumption, potentially threatening food security. Depletion of stratospheric ozone has altered climate in the southern hemisphere in ways that have increased oceanic productivity and consequently the growth, survival and reproduction of many sea birds and mammals. In contrast, warmer sea surface temperatures related to these climate shifts are also correlated with declines in both kelp beds in Tasmania and corals in Brazil. This assessment demonstrates that knowledge of the interactive effects of ozone depletion, UV radiation, and climate change factors on aquatic ecosystems has advanced considerably over the past four years and confirms the importance of considering synergies between environmental factors.

Does allochthony in lakes change across an elevation gradient?.

Ecosystems are subsidized with inputs of mass and energy from their surroundings. These allochthonous inputs regulate many ecosystem characteristics. In inland waters, terrestrial inputs of organic matter regulate food-web structure, ecosystem metabolism, water clarity, and thermal stratification. Future changes in allochthony may be especially pronounced in high-elevation ecosystems due to increases in vegetation and precipitation associated with climate change. Several techniques exist to characterize the degree of allochthony of organic matter in aquatic systems, including metrics such as ΔH, the net isotopic discrimination between water and particulate organic matter (POM) of deuterium stable isotopes, and the fluorescence index (FI), which characterizes the fluorescence of dissolved organic matter (DOM). Despite the importance of allochthonous organic carbon inputs, little is known about either how allochthony varies across elevation gradients or whether different metrics are similarly related to allochthony. We measured AH, FI, and a suite of related water-quality characteristics in 30 lakes across a montane to alpine elevation gradient (2340 to 3205 m) in the Beartooth Mountains of Montana and Wyoming, USA, to understand how FI and AH varied with elevation, with one another, and with other allochthony-related water-quality characteristics. We hypothesized that allochthony of POM and DOM would decrease at higher elevations, with alpine lakes above treeline being more autochthonous compared with low-elevation lakes below treeline. We observed a significant inverse linear relationship between AH and Fl, with both metrics indicating a decrease in allochthony at higher elevations. Characteristics including the natural log of the ratio of concentrations of dissolved organic carbon to chlorophyll a (ln(DOC: Chl)), the spectral slope ratio between different spectra of two wavebands (SR, ratio of spectra at 275-295 to 350-400 nm), and a ratio of diffuse attenuation coefficients at 320 and 380 nm (KR, Kd320: Kd380) varied with both ΔH and FI while pH varied only with ΔH. High-elevation systems were characterized by low ln(DOC: Chl) and K(R), and high S(R) and pH. These results indicate that high-elevation lakes are more autochthonous than low-elevation lakes. The relationships among ΔH, FI, elevation, and other water-quality characteristics provide important insights to understand future changes in carbon cycling in mountain ecosystems.

Effects of UV radiation on aquatic ecosystems and interactions with other environmental factors.

Interactions between climate change and UV radiation are having strong effects on aquatic ecosystems due to feedback between temperature, UV radiation, and greenhouse gas concentration. Higher air temperatures and incoming solar radiation are increasing the surface water temperatures of lakes and oceans, with many large lakes warming at twice the rate of regional air temperatures. Warmer oceans are changing habitats and the species composition of many marine ecosystems. For some, such as corals, the temperatures may become too high. Temperature differences between surface and deep waters are becoming greater. This increase in thermal stratification makes the surface layers shallower and leads to stronger barriers to upward mixing of nutrients necessary for photosynthesis. This also results in exposure to higher levels of UV radiation of surface-dwelling organisms. In polar and alpine regions decreases in the duration and amount of snow and ice cover on lakes and oceans are also increasing exposure to UV radiation. In contrast, in lakes and coastal oceans the concentration and colour of UV-absorbing dissolved organic matter (DOM) from terrestrial ecosystems is increasing with greater runoff from higher precipitation and more frequent extreme storms. DOM thus creates a refuge from UV radiation that can enable UV-sensitive species to become established. At the same time, decreased UV radiation in such surface waters reduces the capacity of solar UV radiation to inactivate viruses and other pathogens and parasites, and increases the difficulty and price of purifying drinking water for municipal supplies. Solar UV radiation breaks down the DOM, making it more available for microbial processing, resulting in the release of greenhouse gases into the atmosphere. In addition to screening solar irradiance, DOM, when sunlit in surface water, can lead to the formation of reactive oxygen species (ROS). Increases in carbon dioxide are in turn acidifying the oceans and inhibiting the ability of many marine organisms to form UV-absorbing exoskeletons. Many aquatic organisms use adaptive strategies to mitigate the effects of solar UV-B radiation (280-315 nm), including vertical migration, crust formation, synthesis of UV-absorbing substances, and enzymatic and non-enzymatic quenching of ROS. Whether or not genetic adaptation to changes in the abiotic factors plays a role in mitigating stress and damage has not been determined. This assessment addresses how our knowledge of the interactive effects of UV radiation and climate change factors on aquatic ecosystems has advanced in the past four years.

Indicators of the effects of climate change on freshwater ecosystems.

Freshwater ecosystems, including lakes, streams, and wetlands, are responsive to climate change and other natural and anthropogenic stresses. These ecosystems are frequently hydrologically and ecologically connected with one another and their surrounding landscapes, thereby integrating changes throughout their watersheds. The responses of any given freshwater ecosystem to climate change depend on the magnitude of climate forcing, interactions with other anthropogenic and natural changes, and the characteristics of the ecosystem itself. Therefore, the magnitude and manner in which freshwater ecosystems respond to climate change is difficult to predict a priori. We present a conceptual model to elucidate how freshwater ecosystems are altered by climate change. We identify eleven indicators that describe the response of freshwater ecosystems to climate change, discuss their potential value and limitations, and describe supporting measurements. Indicators are organized in three inter-related categories: hydrologic, water quality, and ecosystem structure and function. The indicators are supported by data sets with a wide range of temporal and spatial coverage, and they inform important scientific and management needs. Together, these indicators improve the understanding and management of the effects of climate change on freshwater ecosystems.

Solar radiation decreases parasitism in Daphnia.

Climate change and variation in atmospheric ozone are influencing the intensity of ultraviolet radiation (UVR) reaching ecosystems. Changing UVR regimes, in turn, may alter epidemics of infectious disease. This possibility hinges on the sensitivity of epidemiologically relevant traits of host and parasite to UVR. We address this issue using a planktonic system (a zooplankton host, Daphnia dentifera, and its virulent fungal parasite, Metschnikowia bicuspidata). Controlled laboratory experiments, coupled with in situ field incubations of spores, revealed that quite low levels of UVR (as well as longer wavelength light) sharply reduced the infectivity of fungal spores but did not affect host susceptibility to infection. The parasite's sensitivity to solar radiation may underlie patterns in a lake survey: higher penetration of solar radiation into lakes correlated with smaller epidemics that started later in autumn (as incident sunlight declined). Thus, solar radiation, by diminishing infectivity of the parasite, may potently reduce disease.

Climate-induced changes in lake ecosystem structure inferred from coupled neo- and paleoecological approaches.

Over the 20th century, surface water temperatures have increased in many lake ecosystems around the world, but long-term trends in the vertical thermal structure of lakes remain unclear, despite the strong control that thermal stratification exerts on the biological response of lakes to climate change. Here we used both neo- and paleoecological approaches to develop a fossil-based inference model for lake mixing depths and thereby refine understanding of lake thermal structure change. We focused on three common planktonic diatom taxa, the distributions of which previous research suggests might be affected by mixing depth. Comparative lake surveys and growth rate experiments revealed that these species respond to lake thermal structure when nitrogen is sufficient, with species optima ranging from shallower to deeper mixing depths. The diatom-based mixing depth model was applied to sedimentary diatom profiles extending back to 1750 AD in two lakes with moderate nitrate concentrations but differing climate settings. Thermal reconstructions were consistent with expected changes, with shallower mixing depths inferred for an alpine lake where treeline has advanced, and deeper mixing depths inferred for a boreal lake where wind strength has increased. The inference model developed here provides a new tool to expand and refine understanding of climate-induced changes in lake ecosystems.

Environmental effects of ozone depletion and its interactions with climate change: progress report, 2011.

The parties to the Montreal Protocol are informed by three panels of experts. One of these is the Environmental Effects Assessment Panel (EEAP), which deals with two focal issues. The first focus is the effects of increased UV radiation on human health, animals, plants, biogeochemistry, air quality, and materials. The second focus is on interactions between UV radiation and global climate change and how these may affect humans and the environment. When considering the effects of climate change, it has become clear that processes resulting in changes in stratospheric ozone are more complex than believed previously. As a result of this, human health and environmental problems will be longer-lasting and more regionally variable. Like the other panels, the EEAP produces a detailed report every four years; the most recent was published in 2010 (Photochem. Photobiol. Sci., 2011, 10, 173-300). In the years in between, the EEAP produces less detailed and shorter progress reports, which highlight and assess the significance of developments in key areas of importance to the parties. The next full quadrennial report will be published in 2014-2015.

Reintegrating Biology Through the Nexus of Energy, Information, and Matter.

Information, energy, and matter are fundamental properties of all levels of biological organization, and life emerges from the continuous flux of matter, energy, and information. This perspective piece defines and explains each of the three pillars of this nexus. We propose that a quantitative characterization of the complex interconversions between matter, energy, and information that comprise this nexus will help us derive biological insights that connect phenomena across different levels of biological organization. We articulate examples from multiple biological scales that highlight how this nexus approach leads to a more complete understanding of the biological system. Metrics of energy, information, and matter can provide a common currency that helps link phenomena across levels of biological organization. The propagation of energy and information through levels of biological organization can result in emergent properties and system-wide changes that impact other hierarchical levels. Deeper consideration of measured imbalances in energy, information, and matter can help researchers identify key factors that influence system function at one scale, highlighting avenues to link phenomena across levels of biological organization and develop predictive models of biological systems.

Smoke from regional wildfires alters lake ecology.

Wildfire smoke often covers areas larger than the burned area, yet the impacts of smoke on nearby aquatic ecosystems are understudied. In the summer of 2018, wildfire smoke covered Castle Lake (California, USA) for 55 days. We quantified the influence of smoke on the lake by comparing the physics, chemistry, productivity, and animal ecology in the prior four years (2014-2017) to the smoke year (2018). Smoke reduced incident ultraviolet-B (UV-B) radiation by 31% and photosynthetically active radiation (PAR) by 11%. Similarly, underwater UV-B and PAR decreased by 65 and 44%, respectively, and lake heat content decreased by 7%. While the nutrient limitation of primary production did not change, shallow production in the offshore habitat increased by 109%, likely due to a release from photoinhibition. In contrast, deep-water, primary production decreased and the deep-water peak in chlorophyll a did not develop, likely due to reduced PAR. Despite the structural changes in primary production, light, and temperature, we observed little significant change in zooplankton biomass, community composition, or migration pattern. Trout were absent from the littoral-benthic habitat during the smoke period. The duration and intensity of smoke influences light regimes, heat content, and productivity, with differing responses to consumers.

Global data set of long-term summertime vertical temperature profiles in 153 lakes.

Climate change and other anthropogenic stressors have led to long-term changes in the thermal structure, including surface temperatures, deepwater temperatures, and vertical thermal gradients, in many lakes around the world. Though many studies highlight warming of surface water temperatures in lakes worldwide, less is known about long-term trends in full vertical thermal structure and deepwater temperatures, which have been changing less consistently in both direction and magnitude. Here, we present a globally-expansive data set of summertime in-situ vertical temperature profiles from 153 lakes, with one time series beginning as early as 1894. We also compiled lake geographic, morphometric, and water quality variables that can influence vertical thermal structure through a variety of potential mechanisms in these lakes. These long-term time series of vertical temperature profiles and corresponding lake characteristics serve as valuable data to help understand changes and drivers of lake thermal structure in a time of rapid global and ecological change.

Ultraviolet radiation affects invasibility of lake ecosystems by warm-water fish.

Predicting where species invasions will occur remains a substantial challenge in ecology, but identifying factors that ultimately constrain the distribution of potential invaders could facilitate successful prediction. Whereas ultraviolet radiation (UVR) is recognized as an important factor controlling species distribution and community composition, the role of UVR in a habitat invasibility context has not been explored. Here we examine how underwater UVR can regulate warm-water fish invasion. In Lake Tahoe, California and Nevada, USA, established populations of exotic bluegill sunfish (Lepomis macrochirus) are currently limited to turbid, low-UVR embayments. An in situ incubation experiment that manipulated incident UVR exposure of larval bluegill, combined with an assessment of UVR exposure levels in nearshore habitats around Lake Tahoe, demonstrates that UVR can mediate habitat invasibility. Our findings suggest that the susceptibility to invasion by UVR sensitive species may increase in transparent aquatic systems threatened by declining water quality, and they highlight the importance of abiotic factors as regulators of invasion risk in ecosystems.

Correction: Dissolved organic matter protects mosquito larvae from damaging solar UV radiation.

[This corrects the article DOI: 10.1371/journal.pone.0240261.].

Dissolved organic matter protects mosquito larvae from damaging solar UV radiation.

Mosquitoes have increased in their abundance and geographic distribution in northeastern North America, coinciding with an increase in extreme precipitation events and up to a doubling of dissolved organic matter (DOM) concentrations in some inland waters. Increases in DOM can reduce exposure of mosquito larvae to solar ultraviolet (UV) radiation. Although mosquito larvae are most common in shaded habitats, almost nothing is known about their susceptibility to damage by solar UV radiation, or the ability of DOM to create a refuge from damaging UV in their shallow-water habitats. We hypothesize that 1) exposure to solar UV radiation is lethal to mosquito larvae, 2) larvae lack photo-enzymatic repair to fix UV-damaged DNA, and 3) DOM shades larvae from lethal solar UV radiation. We tested these hypotheses with experiments that manipulated UV radiation, the photo-repair radiation necessary for photo-enzymatic DNA repair, and DOM. Exposure to solar UV radiation significantly decreased larval survivorship, while DOM significantly increased it. There was no evidence of photo-enzymatic DNA repair. Our findings confirm that solar UV radiation decreases habitat suitability for mosquito larvae, but DOM provides a refuge from UV. This highlights the need for vector control managers to prioritize high DOM and shaded habitats in their efforts to reduce mosquito populations.

Shedding light on environmentally transmitted parasites: lighter conditions within lakes restrict epidemic size.

Parasite fitness depends on a successful journey from one host to another. For parasites that are transmitted environmentally, abiotic conditions might modulate the success of this journey. Here we evaluate how light, a key abiotic factor, influences spatiotemporal patterns of zooplankton disease where light varies seasonally, across lakes, and with depth in a lake. In an in situ experiment using those three sources of variation, we tested sensitivity of spores of two parasites to ambient light. Infectivity of both parasites was lower when exposed to ambient light in comparison to parasites exposed to otherwise similar conditions in the dark. The more sensitive parasite (the fungus, Metschnikowia) was damaged even under lower ambient light during late fall (November). With this differential sensitivity established, we evaluated links between light environment and natural outbreaks in lakes. Consistent with the incubations, epidemics of the less sensitive parasite (the bacterium, Pasteuria) started earlier in the fall (under higher ambient light), and both parasites had smaller outbreaks in more transparent lakes. Overall, light environment may impact the timing and size of disease outbreaks. Outbreaks could thus become exacerbated by human activities that darken waters, including lake browning associated with climate change and eutrophication.

Deeper waters are changing less consistently than surface waters in a global analysis of 102 lakes.

Globally, lake surface water temperatures have warmed rapidly relative to air temperatures, but changes in deepwater temperatures and vertical thermal structure are still largely unknown. We have compiled the most comprehensive data set to date of long-term (1970-2009) summertime vertical temperature profiles in lakes across the world to examine trends and drivers of whole-lake vertical thermal structure. We found significant increases in surface water temperatures across lakes at an average rate of + 0.37 °C decade-1, comparable to changes reported previously for other lakes, and similarly consistent trends of increasing water column stability (+ 0.08 kg m-3 decade-1). In contrast, however, deepwater temperature trends showed little change on average (+ 0.06 °C decade-1), but had high variability across lakes, with trends in individual lakes ranging from - 0.68 °C decade-1 to + 0.65 °C decade-1. The variability in deepwater temperature trends was not explained by trends in either surface water temperatures or thermal stability within lakes, and only 8.4% was explained by lake thermal region or local lake characteristics in a random forest analysis. These findings suggest that external drivers beyond our tested lake characteristics are important in explaining long-term trends in thermal structure, such as local to regional climate patterns or additional external anthropogenic influences.