Productivity denotes the rate of biomass synthesis in ecosystems and is a fundamental characteristic that frames ecosystem function and management. Limitation of productivity by nutrient availability is an established paradigm for lake ecosystems. Here, we assess the relevance of this paradigm for a majority of the world's small, nutrient-poor lakes, with different concentrations of coloured organic matter. By comparing small unproductive lakes along a water colour gradient, we show that coloured terrestrial organic matter controls the key process for new biomass synthesis (the benthic primary production) through its effects on light attenuation. We also show that this translates into effects on production and biomass of higher trophic levels (benthic invertebrates and fish). These results are inconsistent with the idea that nutrient supply primarily controls lake productivity, and we propose that a large share of the world's unproductive lakes, within natural variations of organic carbon and nutrient input, are limited by light and not by nutrients. We anticipate that our result will have implications for understanding lake ecosystem function and responses to environmental change. Catchment export of coloured organic matter is sensitive to short-term natural variability and long-term, large-scale changes, driven by climate and different anthropogenic influences. Consequently, changes in terrestrial carbon cycling will have pronounced effects on most lake ecosystems by mediating changes in light climate and productivity of lakes.

Carbon dioxide (CO 2 ) evasion from streams and rivers to the atmosphere represents a substantial flux in the global carbon cycle 1-3 . The proportions of CO 2 emitted from streams and rivers that come from terrestrially derived CO 2 or from CO 2 produced within freshwater ecosystems through aquatic metabolism are not well quantified. Here we estimated CO 2 emissions from running waters in the contiguous United States, based on freshwater chemical and physical characteristics and modelled gas transfer velocities at 1463 United States Geological Survey monitoring sites. We then assessed CO 2 production from aquatic metabolism, compiled from previously published measurements of net ecosystem production from 187 streams and rivers across the contiguous United States. We find that CO 2 produced by aquatic metabolism contributes about 28% of CO 2 evasion from streams and rivers with flows between 0.0001 and 19,000 m 3 s −1 . We mathematically modelled CO 2 flux from groundwater into running waters along a stream-river continuum to evaluate the relationship between stream size and CO 2 source. Terrestrially derived CO 2 dominates emissions from small streams, and the percentage of CO 2 emissions from aquatic metabolism increases with stream size. We suggest that the relative role of rivers as conduits for terrestrial CO 2 e ux and as reactors mineralizing terrestrial organic carbon is a function of their size and connectivity with landscapes.Inland waters play a central role in the global carbon (C) cycle by transforming, outgassing and storing more than half of the C they receive from terrestrial ecosystems before delivery to oceans 1-3 . Terrestrial C inputs to freshwaters are often of similar magnitude to terrestrial net ecosystem production (NEP; refs 1,2,4). Consequently, ignoring inland waters in landscape C budgets may overestimate terrestrial CO 2 uptake and storage 1,5 . In fact, not accounting for terrestrial C exports to and emissions from freshwaters could bias terrestrial NEP and net ecosystem exchange measurements by 4-60% (refs 6-8). Despite small areal coverage, running waters are hotspots for CO 2 emissions 3,9 , with high rates of outgassing relative to lake and terrestrial ecosystems on an areal basis 3,10,11 . Given their significant role in landscape C transformations, transport and emissions, there is a fundamental need to understand rates and drivers of C cycling in running waters.A mechanistic understanding of the processes regulating CO 2 emissions from streams and rivers is necessary for sound predictions of the present and future role of freshwaters in global C cycling and the climate system. Inland waters are often supersaturated with CO 2 due to inputs of terrestrially derived CO 2 and in situ aquatic mineralization of terrestrial OC (refs 12-15) (hereafter, 'internal production') as well as abiotic CO 2 production (Supplementary Section 1). CO 2 concentrations and emissions from running waters will thus vary with changes in land cover, climate, terrestrial ecosystem processes, land-water c...

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Abstract. The Arctic is a water-rich region, with freshwater systems covering about 16 % of the northern permafrost landscape. Permafrost thaw creates new freshwater ecosystems, while at the same time modifying the existing lakes, streams, and rivers that are impacted by thaw. Here, we describe the current state of knowledge regarding how permafrost thaw affects lentic (still) and lotic (moving) systems, exploring the effects of both thermokarst (thawing and collapse of ice-rich permafrost) and deepening of the active layer (the surface soil layer that thaws and refreezes each year). Within thermokarst, we further differentiate between the effects of thermokarst in lowland areas vs. that on hillslopes. For almost all of the processes that we explore, the effects of thaw vary regionally, and between lake and stream systems. Much of this regional variation is caused by differences in ground ice content, topography, soil type, and permafrost coverage. Together, these modifying factors determine (i) the degree to which permafrost thaw manifests as thermokarst, (ii) whether thermokarst leads to slumping or the formation of thermokarst lakes, and (iii) the manner in which constituent delivery to freshwater systems is altered by thaw. Differences in thaw-enabled constituent delivery can Published by Copernicus Publications on behalf of the European Geosciences Union. J. E. Vonk et al.: Effects of permafrost thaw on Arctic aquatic ecosystemsbe considerable, with these modifying factors determining, for example, the balance between delivery of particulate vs. dissolved constituents, and inorganic vs. organic materials. Changes in the composition of thaw-impacted waters, coupled with changes in lake morphology, can strongly affect the physical and optical properties of thermokarst lakes. The ecology of thaw-impacted lakes and streams is also likely to change; these systems have unique microbiological communities, and show differences in respiration, primary production, and food web structure that are largely driven by differences in sediment, dissolved organic matter, and nutrient delivery. The degree to which thaw enables the delivery of dissolved vs. particulate organic matter, coupled with the composition of that organic matter and the morphology and stratification characteristics of recipient systems will play an important role in determining the balance between the release of organic matter as greenhouse gases (CO 2 and CH 4 ), its burial in sediments, and its loss downstream. The magnitude of thaw impacts on northern aquatic ecosystems is increasing, as is the prevalence of thaw-impacted lakes and streams. There is therefore an urgent need to quantify how permafrost thaw is affecting aquatic ecosystems across diverse Arctic landscapes, and the implications of this change for further climate warming.

Dissolved organic carbon (DOC) concentrations in lakes are changing globally, but little is known about potential ecosystem impacts.We evaluated the relationship between DOC and whole‐lake primary production in arctic and boreal lakes. Both light extinction (inhibits primary production) and nutrient availability (stimulates primary production) are positively and nonlinearly related to DOC concentration. These nonlinearities create a threshold DOC concentration (4.8 mg L−1), below which the DOC‐primary production relationship is positive, and above which the relationship is negative. DOC concentration varies maximally between regions, creating a unimodal relationship between primary production and DOC that emerges at broader scales because arctic lakes largely fall below the threshold DOC concentration, but boreal lakes fall above it. Our analysis suggests that the impact of DOC trends on lake primary production will vary across lakes and regions as a result of contrasting baseline conditions relative to the DOC threshold.

We investigated productivity at the basal trophic level in 15 unproductive lakes in a gradient ranging from clear-water to brown-water (humic) lakes in northern Sweden. Primary production and bacterial production in benthic and pelagic habitats were measured to estimate the variation in energy mobilization from external energy sources (primary production plus bacterial production on allochthonous organic carbon) along the gradient. Clear-water lakes were dominated by autotrophic energy mobilization in the benthic habitat, whereas humic lakes were dominated by heterotrophic energy mobilization in the pelagic habitat. Whole-lake (benthic + pelagic) energy mobilization was negatively correlated to the light-extinction coefficient, which was determined by colored terrestrial organic matter in the lake water. Thus, variation in the concentration of terrestrial organic matter and its light-absorbing characteristics exerts strong control on the magnitude, as well as on the processes and pathways, of energy mobilization in unproductive lakes. We suggest that unproductive lakes in general are sensitive to input of terrestrial organic matter because of its effects on basal energy mobilization in both benthic and pelagic habitats.

We compared the stable carbon isotopic composition (␦ 13 C) of crustacean zooplankton with that of potential carbon sources in 15 lakes in northern Sweden with different dissolved organic carbon (DOC) concentrations (2-9 mg L Ϫ1 ) to test the hypothesis that zooplankton depended more on allochthonous carbon in humic lakes than in clear-water lakes. Based on ␦ 13 C signature, we found that the pool of organic matter in the lakes was dominated by carbon of allochthonous origin over the whole DOC gradient. Zooplankton were generally depleted in 13 C compared to organic matter in the catchment, particulate organic matter in the lake water, and shallow surface sediment. However, the isotopic composition of zooplankton could not be explained without a significant contribution from both allochthonous and autochthonous carbon sources in all lakes. The relative importance of these two carbon sources did not relate to the concentration of, or proportion between, allochthonous and autochthonous organic carbon in the water. Instead, the proportion between allochthonous and autochthonous carbon in the crustacean zooplankton was consistent with a rather conservative use of the energy mobilized by bacterioplankton and phytoplankton in the lakes.

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.