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Metabolic stoichiometry predicts that dissolved oxygen (O 2) and carbon dioxide (CO 2) in aquatic ecosystems should covary inversely; however, field observations often diverge from theoretical expectations. Here, we propose a suite of metrics describing this O 2 and CO 2 decoupling and introduce a conceptual framework for interpreting these metrics within aquatic ecosystems. Within this framework, we interpret cross-system patterns of high-frequency O 2 and CO 2 measurements in 11 northern lakes and extract emergent insights into the metabolic behavior and the simultaneous roles of chemical and physical forcing in shaping ecosystem processes. This approach leverages the power of high-frequency paired O 2-CO 2 measurements, and yields a novel, integrative aquatic system typology which can also be applicable more broadly to streams and rivers, wetlands and marine systems. Dissolved oxygen (O 2) remains one of the most studied attributes of aquatic ecosystems since the beginning of modern ecology. In 1957, G. E. Hutchinson famously wrote "A skillful limnologist can probably learn more about the nature of a lake from a series of oxygen determinations than from any other kind of chemical data" (Hutchinson 1957). The value of oxygen as an indicator of ecosystem function stems from its role in biogeochemical reactions, where it regulates

Understanding the mechanisms driving carbon dioxide (CO2) concentrations in inland waters is important to foresee CO(2)responses to environmental change, yet knowledge gaps persist regarding which processes are the key drivers. Here we investigated possible drivers across 13 Swedish lakes and streams where the partial pressure of CO2(pCO(2)) has increased over a 21-year period. Overall, we could not identify a single dominating mechanism responsible for the observedpCO(2)increase. In the 8 lakes, we found thatpCO(2)increased, driven either by a possible dissolved organic carbon (DOC) stimulation of microbial mineralization or by water color primary production suppression. In streams, the dominating mechanism for apCO(2)increase was either a change in the carbonate system distribution or a possible nutrient-driven decrease in primary production. This is the first study to demonstrate and explain consistent positivepCO(2)temporal trends in freshwater ecosystems, and our results should be taken into account when predicting future emission of CO(2)from inland waters.

People extensively use lakes and rivers covered by seasonal ice. Although ice cover duration has been declining over the past 150 years for Northern Hemisphere freshwaters, we know relatively little about how ice loss directly affects humans. Here, we synthesize the cultural ecosystem services (i.e., services that provide intangible or nonmaterial benefits) and associated benefits supported by inland ice. We also provide, for the first time, empirical examples that give quantitative evidence for a winter warming effect on a wide range of ice‐related cultural ecosystem services and benefits. We show that in recent decades, warmer air temperatures delayed the opening date of winter ice roads and led to cancellations of spiritual ceremonies, outdoor ice skating races, and ice fishing tournaments. Additionally, our synthesis effort suggests unexploited data sets that allow for the use of integrative approaches to evaluate the interplay between inland ice loss and society. published

Carbon dioxide (CO2) uptake by phytoplankton can significantly reduce the partial pressure of CO2 (pCO2) in lakes and rivers, and thereby CO2 emissions. Presently, it is not known in which inland waters on Earth a significant pCO2 reduction by phytoplankton is likely. Since detailed, comparable carbon budgets are currently not available for most inland waters, we modified a proxy to assess the pCO2 reduction by phytoplankton, originally developed for boreal lakes, for application on a global scale. Using data from 61 rivers and 125 lakes distributed over five continents, we show that a significant pCO2 reduction by phytoplankton is widespread across the temperate and sub-/tropical region, but absent in the cold regions on Earth. More specifically, we found that a significant pCO2 reduction by phytoplankton might occur in 24% of the lakes in the temperate region, and 39% of the lakes in the sub-/tropical region. We also showed that such a reduction might occur in 21% of the rivers in the temperate region, and 5% of the rivers in the sub-/tropical region. Our results indicate that CO2 uptake by phytoplankton is a relevant flux in regional and global carbon budgets. This highlights the need for more accurate approaches to quantify CO2 uptake by primary producers in inland waters, particularly in the temperate and sub-/tropical region.

In the Arctic, rising seawater temperatures and increasing underwater light caused by reductions in sea ice cover are expected to change the structure of arctic marine communities. Substantial, som ...

Northern ecosystems are experiencing some of the most dramatic impacts of global change on Earth. Rising temperatures, hydrological intensification, changes in atmospheric acid deposition and associated acidification recovery, and changes in vegetative cover are resulting in fundamental changes in terrestrial-aquatic biogeochemical linkages. The effects of global change are readily observed in alterations in the supply of dissolved organic matter (DOM)-the messenger between terrestrial and lake ecosystems-with potentially profound effects on the structure and function of lakes. Northern terrestrial ecosystems contain substantial stores of organic matter and filter or funnel DOM, affecting the timing and magnitude of DOM delivery to surface waters. This terrestrial DOM is processed in streams, rivers, and lakes, ultimately shifting its composition, stoichiometry, and bioavailability. Here, we explore the potential consequences of these global change-driven effects for lake food webs at northern latitudes. Notably, we provide evidence that increased allochthonous DOM supply to lakes is overwhelming increased autochthonous DOM supply that potentially results from earlier ice-out and a longer growing season. Furthermore, we assess the potential implications of this shift for the nutritional quality of autotrophs in terms of their stoichiometry, fatty acid composition, toxin production, and methylmercury concentration, and therefore, contaminant transfer through the food web. We conclude that global change in northern regions leads not only to reduced primary productivity but also to nutritionally poorer lake food webs, with discernible consequences for the trophic web to fish and humans.

Boreal lakes can be ice covered for a substantial portion of the year at which time methane (CH4) can accumulate below ice. The amount of CH4 emitted at ice melt is partially determined by the inte ...

The emissions of carbon dioxide (CO2) from inland waters are substantial on a global scale. Yet, the fundamental question remains open which proportion of these CO2 emissions is induced by sunlight via photochemical mineralization of dissolved organic carbon (DOC), rather than by microbial respiration during DOC decomposition. Also, it is unknown on larger spatial and temporal scales how photochemical mineralization compares to other C fluxes in the inland water C cycle. We combined field and laboratory data with atmospheric radiative transfer modeling to parameterize a photochemical rate model for each day of the year 2009, for 1086 lakes situated between latitudes from 55 to 69°N in Sweden. The sunlight-induced production of dissolved inorganic carbon (DIC) averaged 3.8 ± 0.04 g C m-2 yr-1, which is a flux comparable in size to the organic carbon burial in the lake sediments. Countrywide, 151 ± 1 kt C yr-1 was produced by photochemical mineralization, corresponding to about 12% of total annual mean CO2 emissions from Swedish lakes. With a median depth of 3.2 m, the lakes were generally deep enough that incoming, photochemically active photons were absorbed in the water column. This resulted in a linear positive relationship between DIC photoproduction and the incoming photon flux, which correspond to the absorbed photons. Therefore, the slope of the regression line represents the wavelength- and depth-integrated apparent quantum yield of DIC photoproduction. We used this relationship to obtain a first estimate of DIC photoproduction in lakes and reservoirs worldwide. Global DIC photoproduction amounted to 13 and 35 Mt C yr-1 under overcast and clear sky, respectively. Consequently, these directly sunlight-induced CO2 emissions contribute up to about one tenth to the global CO2 emissions from lakes and reservoirs, corroborating that microbial respiration contributes a substantially larger share than formerly thought, and generate annual C fluxes similar in magnitude to the C burial in natural lake sediments worldwide.

Peer-reviewed. Copyright: © 2016 Woolway et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.PLoS ONE 11(3): e0152466. doi:10.1371/journal.pone.0152466, Ecological and biogeochemical processes in lakes are strongly dependent upon water temperature. Long-term surface warming of many lakes is unequivocal, but little is known about the comparative magnitude of temperature variation at diel timescales, due to a lack of appropriately resolved data. Here we quantify the pattern and magnitude of diel temperature variability of surface waters using high-frequency data from 100 lakes. We show that the near-surface diel temperature range can be substantial in summer relative to long-term change and, for lakes smaller than 3 km2, increases sharply and predictably with decreasing lake area. Most small lakes included in this study experience average summer diel ranges in their near-surface temperatures of between 4 and 7°C. Large diel temperature fluctuations in the majority of lakes undoubtedly influence their structure, function and role in biogeochemical cycles, but the full implications remain largely unexplored., This project was funded by OTT Hydrometry Ltd and a University College London impact award awarded to RIW and is part of the Natural Environment Research Council (NERC) Sensor Network project United Kingdom Lake Ecological Observatory Network (UKLEON;NE/I007407/1). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.

The availability of substrates is one of the most important environmental constraints on the diversity and functioning of microorganisms. Substrate quantity and quality as well as the metabolic features of heterotrophic microorganisms determine the efficiency, speed and type of transformation that can occur in nature. As such their interplay with the environment regulates how much carbon and energy is incorporated by bacteria and subsequently reaches higher trophic levels. In lakes the bulk substrate that is available for bacteria is composed of a complex mixture of compounds, varying in lability and distribution in the environment. This thesis addresses the coupling of organic substrates, their metabolic use and the composition and ecology of the microbial community. Controlled laboratory experiments with mixed bacterial communities in either batch cultures or chemostats were designed to shed further light on bacterial use of labile and quantitatively significant carbon compounds.I show that different amino acid substrates only exert a minor influence on bacterioplankton community composition and growth. Hence the ability to use a wide range of such abundantly produced protein monomers seems to be widespread among freshwater bacteria. In contrast, when acetate was provided as the only carbon substrate, in either pulsed or continuous amendments, this very different substrate input mode had a strong effect on bacterial community composition. Biomass yield, for example, was twice as high when acetate was given in the form of pulses rather than provided continuously.In another set of experiments, I show that the oxidation of the globally significant greenhouse gas methane is a process that can potentially take place at the water-ice interface of seasonally ice-covered lakes and was not constrained by temperature as suggested in previous studies. This work also suggests that methane oxidation in ice-covered lakes can be constrained by competition for nutrients between specialized methanotrophs and heterotrophic bacteria.Combined these studies suggest that some labile substrates cause minor selection on bacterial community structure and functioning. This probably reflects the competitive advantage of using a broad range of low molecular weight substrates. However, as in the case of methanotrophs there is specialization for a specific low molecular weight substrate such as methane. In which case, competition with other community members i.e. for nutrients can constrain methane oxidation. In both cases it might however not depend just on the availability of substrate, but also on how substrates are distributed in time and space.

Bacterioplankton growth is often directly or indirectly controlled by external energy subsidies via organic matter inputs or solar radiation. We carried out a mesocosm experiment to assess how bact ...

Freshwater ice dominates the Arctic terrestrial environment and significantly impacts bio-physical and socio-economic systems. Unlike other major cryospheric components that either blanket large expanses (e.g., snow, permafrost, sea ice) or are concentrated in specific locations, lake and river ice are interwoven into the terrestrial landscape through major flow and storage networks. For instance, the headwaters of large ice-covered rivers extend well beyond the Arctic while many northern lakes owe their genesis to broader cryospheric changes. The effects of freshwater ice on climate mostly occur at the local/regional scale, with the degree of influence dependent on the magnitude, timing, location, and duration of ice cover, and the size of the water body. Freshwater-ice formation, growth, decay, and break-up are influenced by climatic variables that control surface heat fluxes, but these differ markedly between lakes and rivers. Despite the importance of freshwater ice, there has been a recent reduction in observational recordings.

Paleolimnological evidence from some Arctic lakes suggests that longer ice-free seasons have been experienced since the beginning of the nineteenth century. It has been inferred from some additional records that many Arctic lakes may have crossed an important ecological threshold as a result of recent warming. In the instrumental record, long-term trends exhibit increasingly later freeze-ups and earlier break-ups, closely corresponding to increasing air temperature trends, but with greater sensitivity at the more temperate latitudes. Broad spatial patterns in these trends are also related to major atmospheric circulation patterns. Future projections of lake ice indicate increasingly later freeze-ups and earlier break-ups, decreasing ice thickness, and changes in cover composition, particularly white-ice. For rivers, projected future decreases in south to north air-temperature gradients suggest that the severity of ice-jam flooding may be reduced but this could be mitigated by changes in the magnitude of spring snowmelt.

To evaluate climate and atmospheric deposition induced physical and water chemical changes and their effects on phytoplankton communities, we used complete time series (14 years, monthly measurements during the growing season) of 18 physical and chemical variables and phytoplankton data from 13 nutrient-poor Swedish reference lakes along a latitudinal gradient. We found numerous strong significant changes over time that were most coherent among lakes for sulfate concentrations, conductivity, calcium, magnesium, chloride, potassium, water color, surface water temperature and the intensity of thermal stratification. Despite these pronounced coherent physical and water chemical changes over Sweden, the phytoplankton biomass and species richness of six phytoplankton groups, measured at the same time as the water chemical variables, showed only few and weak significant changes over time. The only coherent significant change over Sweden, occurring in seven lakes, was observed in the species richness of chlorophytes. The number of chlorophyte taxa significantly declined over Sweden. Using a partial least square model for each lake, we attributed the decline primarily to an increase in water temperatures and water color, which were among the most important variables for the model performance of each lake. All other taxonomic groups were driven primarily by non-coherent changes in nutrient concentrations, pH and probably also non-coherent grazing pressure. We concluded that coherent phytoplankton responses can only be achieved for taxonomic groups that are driven primarily by coherent physical/chemical changes. According to our study, chlorophytes belong to such a group, making them possible global change indicators. Our findings give new insights into global change effects on different phytoplankton taxonomic groups in nutrient-poor lakes.

While there is a general sense that lakes can act as sentinels of climate change, their efficacy has not been thoroughly analyzed. We identified the key response variables within a lake that act as indicators of the effects of climate change on both the lake and the catchment. These variables reflect a wide range of physical, chemical, and biological responses to climate. However, the efficacy of the different indicators is affected by regional response to climate change, characteristics of the catchment, and lake mixing regimes. Thus, particular indicators or combinations of indicators are more effective for different lake types and geographic regions. The extraction of climate signals can be further complicated by the influence of other environmental changes, such as eutrophication or acidification, and the equivalent reverse phenomena, in addition to other land-use influences. In many cases, however, confounding factors can be addressed through analytical tools such as detrending or filtering. Lakes are effective sentinels for climate change because they are sensitive to climate, respond rapidly to change, and integrate information about changes in the catchment.

Climate change effects on freshwater biogeochemistry and riverine loads of biogenic elements to the Baltic Sea are not straight forward and are difficult to distinguish from other human drivers such as atmospheric deposition, forest and wetland management , eutrophication and hydrological alterations. Eutrophication is by far the most well-known factor affecting the biogeochemistry of the receiving waters in the various sub-basins of the Baltic Sea. However, the present literature review reveals that climate change is a compounding factor for all major drivers of freshwater biogeochemistry discussed here, although evidence is still often based on short-term and/or small-scale studies.

Peer reviewed. ©2015. The Authors.This is an open access article under theterms of the Creative CommonsAttribution-NonCommercial-N oDerivsLicense, which permits use and distri-bution in any medium, provided theoriginal work is properly cited, the use isnon-commerc ial and no modificationsor adaptations are made., In this first worldwide synthesis of in situ and satellite-derived lake data, we find that lake summer surface water temperatures rose rapidly (global mean = 0.34°C decade 1) between 1985 and 2009. Our analyses show that surface water warming rates are dependent on combinations of climate and local characteristics, rather than just lake location, leading to the counterintuitive result that regional consistency in lake warming is the exception, rather than the rule. The most rapidly warming lakes are widely geographically distributed, and their warming is associated with interactions among different climatic factors —from seasonally ice-covered lakes in areas where temperature and solar radiation are increasing while cloud cover is diminishing (0.72°C decade 1) to ice-free lakes experiencing increases in air temperature and solar radiation (0.53°C decade 1). The pervasive and rapid warming observed here signals the urgent need to incorporate climate impacts into vulnerability assessments and adaptation efforts for lakes.

Changes in the timing, composition, and intensity of freshwater phytoplankton blooms are known to have an impact on water quality and aquatic ecosystem functions. Factors provoking these changes are, therefore, of major importance. In Lake Erken in southeastern Sweden considerable changes in the timing and large variations in the composition of phytoplankton spring peaks have been observed during the past 45 yr. Here we show that long-term changes and variations in Lake Erken are strongly related to a single global parameter-the North Atlantic Oscillation (NAO). Even regional parameters that are known to have most influence on the spring development of phytoplankton such as ice break-up and nutrient concentrations could not provide a more conclusive explanation of the observed changes in spring phytoplankton, making the NAO a very powerful and simple tool in determining the timing and composition of phytoplankton spring peaks in a temperate lake.

We performed intensive sediment trap studies at different water depths and locations in nine Swedish lakes to determine the flux, distribution, and origin of settling particulate matter in the pelagic zone. From these data, we developed a dynamic model that includes important mechanisms controlling fluxes of autochthonous, allochthonous, and resuspended particles in lakes during both stratification and turnover. The data suggest that during the period of turnover, sediment resuspension is a function of direct wind/wave activities. During these periods, resuspended particles are quite evenly distributed in the water column. During stratification, however, particles in deeper lakes are enriched in the hypolimnion. We suggest that these particles are resuspended due mainly to internal water motions. The model describes how much and at which water depths material is resuspended, and how this material is distributed in the water column. This includes the process of sediment focusing. The model allows determination of sediment accumulation at different water depths and of the boundary between erosion, transportation, and accumulation of sediment. Furthermore, the model shows the time dependence of the flux and distribution of settling particles in relation to changes in wind speed. The model is driven by wind, stratification, and lake morphometric data that can be determined with good accuracy. Some coefficients describing particle dynamics are also needed, e.g. particle settling velocity, vertical water mixing velocity, and sediment resuspension rate. The most uncertain coefficients were subjected to sensitivity analyses. The model can explain differences over time in the flux, origin, and distribution of settling particles in different lakes. Because it has earlier been recognized that the flux and distribution of settling particles can be of great importance for the dispersal, burial, biouptake, and ecological effects of nutrients and contaminants, the model may provide valuable information for lake management.

Phytoplankton, an assemblage of suspended, primarily autotrophic single cells and colonies, forms part of the base of the pelagic food chain in lakes. The responses of phytoplankton to anthropogenic pressures frequently provide the most visible indication of a long-term change in water quality. Several attributes related to the growth and composition of phytoplankton, such as their community structure, abundance as well as the frequency and the intensity of blooms, are included as indicators of water quality in the Water Framework Directive. The growth and seasonal succession of phytoplankton is regulated by a variety of external as well as internal factors (Reynolds et al., 1993; Reynolds, 2006). Among the most important external factors are light, temperature, and those associated with the supply of nutrients from point and diffuse sources in the catchment. The internal factors include the residence time of the lakes, the underwater light regime and the mixing characteristics of the water column. The schematic diagram (Fig. 14.1) shows some of the ways in which systematic changes in the climate can modulate these seasonal and inter-annual variations. The effects associated with the projected changes in the rainfall are likely to be most pronounced in small lakes with short residence times (see George et al., 2004 for some examples). In contrast, those connected with the projected changes in irradiance and wind mixing, are likely to be most important in deep, thermally stratified lakes.

Recent increases in concentrations of dissolved organic carbon (DOC) in lakes and rivers over large regions have been related to both changes in the climate and in atmospheric deposition chemistry. Using a data set of 1041 boreal lakes along a 13° latitudinal gradient, sampled in 1995, 2000, and 2005, and an additional data set of 90 lakes along a 1000-m altitudinal gradient at 68°N, we show that DOC concentrations increase in a nonlinear way along a latitudinal and altitudinal temperature gradient. The nonlinear relation of DOC to increasing temperatures was consistent over space and time. Out of 14 meteorological, catchment, morphometric, and atmospheric deposition variables tested, the variable best explaining this kind of nonlinear pattern was the number of days when air temperatures exceeded 0°C, i.e., the duration of the main growing and runoff season (DT>0). Using DT>0 as an input variable, we were able to predict the nonlinear temperature response of DOC concentrations, both spatially (R2 = 0.90, p < 0.0001) and temporally (R2 = 0.90, p < 0.0001). DT>0 has an advantage over other variables because it includes the time factor, which is decisive for the duration that biogeochemical processes can take place. We suggest that DOC concentrations in lakes are influenced by climate change and that present temperature increases over Sweden result in an accelerated DOC increase toward warmer geographical regions.

Recent studies have highlighted the impact of the winter North Atlantic Oscillation (NAO) on water temperature, ice conditions, and spring plankton phenology in specific lakes and regions in Europe. Here, we use meta-analysis techniques to test whether 18 lakes in northern, western, and central Europe respond coherently to winter climate forcing, and to assess the persistence of the winter climate signal in physical, chemical, and biological variables during the year. A meta-analysis approach was chosen because we wished to emphasize the overall coherence pattern rather than individual lake responses. A particular strength of our approach is that time-series from each of the 18 lakes were subjected to the same robust statistical analysis covering the same 23-year period. Although the strongest overall coherence in response to the winter NAO was exhibited by lake water temperatures, a strong, coherent response was also exhibited by concentrations of soluble reactive phosphorus and soluble reactive silicate, most likely as a result of the coherent response exhibited by the spring phytoplankton bloom. Lake nitrate concentrations showed significant coherence in winter. With the exception of the cyanobacterial biomass in summer, phytoplankton biomass in all seasons was unrelated to the winter NAO. A strong coherence in the abundance of daphnids during spring can most likely be attributed to coherence in daphnid phenology. A strong coherence in the summer abundance of the cyclopoid copepods may have been related to a coherent change in their emergence from resting stages. We discuss the complex nature of the potential mechanisms that drive the observed changes.

We determined relative nitrate-nitrogen (NO3- N) loss rates in 100 north-mid-European lakes from late spring to summer by using the exponential function N-2 5 N-1e(-k)( (t)(2) - (t)(2)), where N-1 and N-2 are NO3- N concentrations at the beginning (t(1)) and the end (t(2)) of the time interval, respectively, and k is the specific NO3- N loss rate. We found that k decreased with increasing lake depth. Adjusting k to the lake depth (k(adj)), we observed that k(adj) was positively related to spring NO3-N concentrations, but this relationship became insignificant at mean lake depths exceeding 12.5 m. A relationship between k(adj) and spring NO3- N concentrations in lakes shallower than 12.5 m implies that changes in spring NO3-N concentrations influence the NO3- N loss rate and thereby summer NO3- N concentrations. Time series from one Estonian, one German, and 14 Swedish lakes shallower than 12.5 m since 1988 revealed that May to August NO3-N concentrations have decreased over time everywhere, and the number of time periods exhibiting a NO3-N depleted condition, i.e., NO3-N levels below 10 mu g L-1, in these lakes has tripled since 1988. We explained the decreasing NO3-N concentrations by a reduction in external nitrogen loading including atmospheric deposition, and by changes in climate. The observed prolongation of NO3- N depleted conditions might be one possible explanation for the increasing occurrence of nitrogen- fixing cyanobacteria in a variety of lake ecosystems.

The North Atlantic Oscillation (NAO) is commonly presented as an easy and reliable index that can be used to study climatic effects on aquatic ecosystems. Here, the NAO winter index (NAO(w)) was applied to determine effects of winter climate changes on 13 water chemical variables measured monthly at 16 sites in Sweden's three largest lakes (Vanern, Vattern, and Malaren). The NAO(w) strongly affected meteorological, physical, and consequently also chemical conditions. Significant relationships were numerous in Sweden's three largest lakes, but they exhibited little agreement among lakes and even across a lake. Synchronous relationships between the NAO(w) and water chemistry among lake sites were restricted to variables closely linked to surface-water temperature (i.e., reactive silica and pH in May). The weaker the association of a lake variable with water temperature, the weaker the mean NAO(w) signal on the variable over the 16 lake sites. The results of this study might facilitate the prediction of lake ecosystem responses to future changes in the weather over a large region.

The North Atlantic Oscillation (NAO) affects the physics, hydrology, chemistry and biology of freshwater ecosystems over a large part of the Northern Hemis- phere. Physical impacts of the NAO include effects on lake temperature profiles, lake ice phenology, river runoff and lake water levels. These physical and hydrological responses influence the chemistry and biology of fresh waters by affecting the leaching of nutrients from the soil and by altering the distribution of nutrients and oxygen in lakes. Finally, the population dynamics of freshwater organisms on several trophic levels including autotrophs, herbivores and vertebrate predators are directly and indirectly linked to the NAO via food-web interactions. As a result, the effects of mild winters associated with the positive index phase of the NAO can influence the food-web characteristics of lakes in summer. A considerable body of evidence documents the importance of these indirect and food-web mediated effects of the NAO, which might even result in ecosystem regime shifts. Owing to the large-scale impact of the NAO, lakes exhibit spatial coherence over large areas with respect to both physical and biological properties. This coherence is modified by geographical factors such as altitude and latitude, and by lake-specific characteristics such as depth and trophic status.

Winters in Sweden have become warmer in the 1990s, and as a consequence the timing of ice break-up and the growth and decline of spring phytoplankton has shifted, starting earlier. Even spring temperatures have become warmer, leading to an earlier beginning of the summer phytoplankton growth. The spring-ward shift in phytoplankton population growth has resulted in an extension of the growing season by at least one month. Although mean total phytoplankton biomass from May to October has not increased, the spring and early summer biomass of temperature-sensitive phytoplankton groups, such as cyanobacteria and chlorophytes, has increased in the 1990s. No increase was noted for other phytoplankton groups. Considering that some species of cyanobacteria that commonly occur during a summer bloom, such as Anabaena, Aphanizomenon, and Microcystis, can be toxic, the effect of warmer winters on aquatic ecosystems is potentially far-reaching.

The North Atlantic Oscillation (NAO) is a regional weather pattern that can be quantified by differences in sea-level pressure measured at stations close to the centres of the Azores High and the Iceland Low (HURRELL 1995). Recent studies show that, especially in winter, changes in the temperature regime (e.g. PLAUDT et al. 1995), snowfall (BENISTON 1997), west wind stress (ROGERS 1997) and the timing of lake ice break-up (LIVINGSTONE 2000) are related to the NAO in countries surrounding the North Atlantic. In this study, phytoplankton has been chosen to demonstrate the effects of a change in the NAO, and thereby changes in the winter climate, on pelagic ecosystems. The phytoplankton community is an ideal sentinel of changes in climatic conditions because it is dominated by short-lived organisms that respond rapidly to changes in the weather (CATALAN & FEE 1994). It is hypothesised that a change in the NAO has an effect on phytoplankton that is similar in time and magnitude in European lakes.

A strong relationship (r > 0.99) between settling particulate inorganic matter (SPIM) and total settling particulate matter (SPM) was observed in 315 samples from sedimentation traps in two Swedish lakes, Lake Erken and Lake Limmaren. This relationship can be used to distinguish between different sources of SPM in pelagic systems when the inflow of allochthonous particulate matter is negligible. It is possible to quantify the proportion of material from planktonic production as well as from sediment resuspension in the flux of settling or suspended particulate matter. Furthermore, fluxes of resuspended organic particles can be quantified and distinguished from fluxes of planktonic organic particles. Although Lake Erken and Lake Limmaren are quite different with respect to trophic level, lake surface area and water depth, the estimated proportion of resuspended particles in sedimentation traps was similar in both lakes. Resuspended particulate matter ranged from 35% of the total settling particulate matter up to 99%, with annual means of 83-94%. Moreover, for both lakes it was estimated that, even in epilimnetic traps, only 2046% (annual means) of the total organic settling particulate matter was from planktonic production, whereas 54-80% (annual means) was from sediment resuspension. This is likely to have important consequences for fluxes of nutrients and contaminants.

We explore the role of lakes in carbon cycling and global climate, examine the mechanisms influencing carbon pools and transformations in lakes, and discuss how the metabolism of carbon in the inland waters is likely to change in response to climate. Furthermore, we project changes as global climate change in the abundance and spatial distribution of lakes in the biosphere, and we revise the estimate for the global extent of carbon transformation in inland waters. This synthesis demonstrates that the global annual emissions of carbon dioxide from inland waters to the atmosphere are similar in magnitude to the carbon dioxide uptake by the oceans and that the global burial of organic carbon in inland water sediments exceeds organic carbon sequestration on the ocean floor. The role of inland waters in global carbon cycling and climate forcing may be changed by human activities, including construction of impoundments, which accumulate large amounts of carbon in sediments and emit large amounts of methane to the atmosphere. Methane emissions are also expected from lakes on melting permafrost. The synthesis presented here indicates that (1) inland waters constitute a significant component of the global carbon cycle, (2) their contribution to this cycle has significantly changed as a result of human activities, and (3) they will continue to change in response to future climate change causing decreased as well as increased abundance of lakes as well as increases in the number of aquatic impoundments.