In-Lake Processes Offset Increased Terrestrial Inputs of Dissolved Organic Carbon and Color to Lakes

Increased color in surface waters, or browning, can alter lake ecological function, lake thermal stratification and pose difficulties for drinking water treatment. Mechanisms suggested to cause browning include increased dissolved organic carbon (DOC) and iron concentrations, as well as a shift to more colored DOC. While browning of surface waters is widespread and well documented, little is known about why some lakes resist it. Here, we present a comprehensive study of Mälaren, the third largest lake in Sweden. In Mälaren, the vast majority of water and DOC enters a western lake basin, and after approximately 2.8 years, drains from an eastern basin. Despite 40 years of increased terrestrial inputs of colored substances to western lake basins, the eastern basin has resisted browning over this time period. Here we find the half-life of iron was far shorter (0.6 years) than colored organic matter (A₄₂₀ ; 1.7 years) and DOC as a whole (6.1 years). We found changes in filtered iron concentrations relate strongly to the observed loss of color in the western basins. In addition, we observed a substantial shift from colored DOC of terrestrial origin, to less colored autochthonous sources, with a substantial decrease in aromaticity (-17%) across the lake. We suggest that rapid losses of iron and colored DOC caused the limited browning observed in eastern lake basins. Across a wider dataset of 69 Swedish lakes, we observed greatest browning in acidic lakes with shorter retention times (< 1.5 years). These findings suggest that water residence time, along with iron, pH and colored DOC may be of central importance when modeling and projecting changes in brownification on broader spatial scales.     

Nitrogen effects on the pelagic food web are modified by dissolved organic carbon

Global environmental change has altered the nitrogen (N) cycle and enhanced terrestrial dissolved organic carbon (DOC) loadings to northern boreal lakes. However, it is still unclear how enhanced N availability affects pelagic food web efficiency (FWE) and crustacean zooplankton growth in N limited boreal lakes. Here, we performed in situ mesocosm experiments in six unproductive boreal Swedish lakes, paired across a DOC gradient, with one lake in each pair fertilized with N (2011: reference year; 2012, 2013: impact years). We assessed how zooplankton growth and FWE were affected by changes in pelagic energy mobilization (PEM), food chain length (phytoplankton versus bacterial production based food chain, i.e. PP:BP), and food quality (seston stoichiometry) in response to N fertilization. Although PP, PEM and PP:BP increased in low and medium DOC lakes after N fertilization, consumer growth and FWE were reduced, especially at low DOC—potentially due to reduced phytoplankton food quality [increased C: phosphorus (P); N:P]. At high DOC, N fertilization caused modest increases in PP and PEM, with marginal changes in PP:BP and phytoplankton food quality, which, combined, led to a slight increase in zooplankton growth and FWE. Consequently, at low DOC (<12 mg L^?1), increased N availability lowers FWE due to mismatches in food quality demand and supply, whereas at high DOC this mismatch does not occur, and zooplankton production and FWE may increase. We conclude that the lake DOC level is critical for predicting the effects of enhanced inorganic N availability on pelagic productivity in boreal lakes.     

Browning reduces the availability—but not the transfer—of essential fatty acids in temperate lakes

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A Framework for Understanding Variation in Pelagic Gross Primary Production of Lake Ecosystems

Light and nutrient availability are key physiological constraints for primary production. Widespread environmental changes are causing variability in loads of terrestrial dissolved organic carbon (DOC) and nutrients from watersheds to lakes, contributing to simultaneous changes in both light and nutrient supply. Experimental evidence highlights the potential for these watershed loads to create complex and context-dependent responses of within-lake primary production; however, the field lacks a predictive model to investigate these responses. We embedded a well-established physiological model of phytoplankton growth within an ecosystem model of nutrient and DOC supply to assess how simultaneous changes in DOC and nutrient loads could impact pelagic primary production in lakes. The model generated a unimodal relationship between GPP and DOC concentration when loads of DOC and nutrients were tightly correlated across space or time. In this unimodal relationship, the magnitude of the peak GPP was primarily determined by the DOC-to-nutrient ratio of the load, and the location of the peak along the DOC axis was primarily determined by lake area. Greater nutrient supply relative to DOC load contributed to greater productivity, and larger lake area increased light limitation for primary producers at a given DOC concentration, owing to the positive relationship between lake area and epilimnion depth. When loads of DOC and nutrients were not tightly correlated in space or time, the model generated a wedge-shaped pattern between GPP and DOC, consistent with spatial surveys from a global set of lakes. Our model is thus capable of unifying the diversity of empirically observed spatial and temporal responses of lake productivity to DOC and mineral nutrient supply presented in the literature, and provides qualitative predictions for how lake pelagic primary productivity may respond to widespread environmental changes.     

Effects of humic stress on the zooplankton from clear and DOC‐rich lakes

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Bottom‐up and top‐down effects of browning and warming on shallow lake food webs

Productivity and trophic structure of aquatic ecosystems result from a complex interplay of bottom‐up and top‐down forces that operate across benthic and pelagic food web compartments. Projected global changes urge the question how this interplay will be affected by browning (increasing input of terrestrial dissolved organic matter), nutrient enrichment and warming. We explored this with a process‐based model of a shallow lake food web consisting of benthic and pelagic components (abiotic resources, primary producers, grazers, carnivores), and compared model expectations with the results of a browning and warming experiment in nutrient‐poor ponds harboring a boreal lake community. Under low nutrient conditions, the model makes three major predictions. (a) Browning reduces light and increases nutrient supply; this decreases benthic and increases pelagic production, gradually shifting productivity from the benthic to the pelagic habitat. (b) Because of active habitat choice, fish exert top‐down control on grazers and benefit primary producers primarily in the more productive of the two habitats. (c) Warming relaxes top‐down control of grazers by fish and decreases primary producer biomass, but effects of warming are generally small compared to effects of browning and nutrient supply. Experimental results were consistent with most model predictions for browning: light penetration, benthic algal production, and zoobenthos biomass decreased, and pelagic nutrients and pelagic algal production increased with browning. Also consistent with expectations, warming had negative effects on benthic and pelagic algal biomass and weak effects on algal production and zoobenthos and zooplankton biomass. Inconsistent with expectations, browning had no effect on zooplankton and warming effects on fish depended on browning. The model is applicable also to nutrient‐rich systems, and we propose that it is a useful tool for the exploration of the consequences of different climate change scenarios for productivity and food web dynamics in shallow lakes, the worldwide most common lake type.     

Life history constraints explain negative relationship between fish productivity and dissolved organic carbon in lakes

Resource availability constrains the life history strategies available to organisms and may thereby limit population growth rates and productivity. We used this conceptual framework to explore the mechanisms driving recently reported negative relationships between fish productivity and dissolved organic carbon (DOC) concentrations in lakes. We studied populations of bluegill (Lepomis macrochirus) in a set of lakes with DOC concentrations ranging from 3 to 24 mg/L; previous work has demonstrated that primary and secondary productivity of food webs is negatively related to DOC concentration across this gradient. For each population, we quantified individual growth rate, age at maturity, age‐specific fecundity, maximum age, length‐weight and length‐egg size relationships, and other life history characteristics. We observed a strong negative relationship between maximum size and DOC concentration; for instance, fish reached masses of 150 to 260 g in low‐DOC lakes but <120 g in high‐DOC lakes. Relationships between fecundity and length, and between egg size and length, were constant across the DOC gradient. Because fish in high‐DOC lakes reached smaller sizes but had similar fecundity and egg size at a given size, their total lifetime fecundity was as much as two orders of magnitude lower than fish in low‐DOC lakes. High DOC concentrations appeared to constrain the range of bluegill life history strategies available; populations in high‐DOC lakes always had low initial growth rates and high ages at maturity, whereas populations in low‐DOC showed higher variability in these traits. This was also the case for the intrinsic rates of natural increase of these populations, which were always low at the high end of the DOC gradient. The potentially lower capacity for fish populations in high‐DOC lakes to recover from exploitation has clear implications for the sustainable management of recreational fisheries in the face of considerable spatial heterogeneity and ongoing temporal change in lake DOC concentrations.     

Net pelagic heterotrophy in mesotrophic and oligotrophic basins of a large, temperate lake

Understanding the effects of trophic status and dissolved organic carbon concentration (DOC) on lake carbon cycling is essential for accurate ecosystem carbon models. Using isotopically labelled substrates we assessed spatial and temporal variability in bacterial respiration (BR) and algal primary production (PP) in two trophically, morphometrically and hydrologically different basins in Loch Lomond, a large temperate lake in Scotland. GIS modelling was used to construct a whole lake balance for bacterial production/respiration and PP, and from this the proportion of heterotrophy fuelled by allochthonous carbon was estimated. We tested the hypotheses that trophic status and DOC concentration affect the balance between PP and BR and examined which is the more significant driving factor. Additionally we estimated the percentage of BR that is fuelled by terrestrial carbon. PP varied seasonally and showed inter-basin homogeneity. BR was greatest in the mesotrophic south basin in autumn, which corresponded to measured peak DOC input, though over an annual cycle no relationship was observed between BR and DOC concentration. The PP:BR ratio was 0.37 ± 0.30 and 0.3 ± 0.45 in the north and south basins, respectively, assuming a bacterial growth efficiency of 0.1. We have found that allochthonous carbon potentially supports a substantial quantity of pelagic production, even during periods of high photosynthesis. Less productive systems are thought to be dominated by heterotrophic processes. However, we have found that the mesotrophic basin of a large lake to be as heterotrophic as its neighbouring oligotrophic basin, an observation that has implications for our understanding of modelling of the role of lakes in linking the terrestrial-atmospheric carbon cycle.     

Climate and landscape influence on indicators of lake carbon cycling through spatial patterns in dissolved organic carbon

Freshwater ecosystems are strongly influenced by both climate and the surrounding landscape, yet the specific pathways connecting climatic and landscape drivers to the functioning of lake ecosystems are poorly understood. Here, we hypothesize that the links that exist between spatial patterns in climate and landscape properties and the spatial variation in lake carbon (C) cycling at regional scales are at least partly mediated by the movement of terrestrial dissolved organic carbon (DOC) in the aquatic component of the landscape. We assembled a set of indicators of lake C cycling (bacterial respiration and production, chlorophyll a, production to respiration ratio, and partial pressure of CO₂), DOC concentration and composition, and landscape and climate characteristics for 239 temperate and boreal lakes spanning large environmental and geographic gradients across seven regions. There were various degrees of spatial structure in climate and landscape features that were coherent with the regionally structured patterns observed in lake DOC and indicators of C cycling. These different regions aligned well, albeit nonlinearly along a mean annual temperature gradient; whereas there was a considerable statistical effect of climate and landscape properties on lake C cycling, the direct effect was small and the overall effect was almost entirely overlapping with that of DOC concentration and composition. Our results suggest that key climatic and landscape signals are conveyed to lakes in part via the movement of terrestrial DOC to lakes and that DOC acts both as a driver of lake C cycling and as a proxy for other external signals.     

Light and nutrient control phytoplankton biomass responses to global change in northern lakes

Global change affects terrestrial loadings of colored dissolved organic carbon (DOC) and nutrients to northern lakes. Still, little is known about how phytoplankton respond to changes in light and nutrient availability across gradients in lake DOC. In this study, we used results from whole‐lake studies in northern Sweden to show that annual mean phytoplankton biomass expressed unimodal curved relationships across lake DOC gradients, peaking at threshold DOC levels of around 11 mg/L. Whole‐lake single nutrient enrichment in selected lakes caused elevated biomass, with most pronounced effect at the threshold DOC level. These patterns give support to the suggested dual control by DOC on phytoplankton via nutrient (positively) and light (negatively) availability and imply that the lakes' location along the DOC axis is critical in determining to what extent phytoplankton respond to changes in DOC and/or nutrient loadings. By using data from the large Swedish Lake Monitoring Survey, we further estimated that 80% of northern Swedish lakes are below the DOC threshold, potentially experiencing increased phytoplankton biomass with browning alone, and/or combined with nutrient enrichment. The results support the previous model results on effects of browning and eutrophication on lake phytoplankton, and provide important understanding of how northern lakes may respond to future global changes.     

Whole-Lake Sugar Addition Demonstrates Trophic Transfer of Dissolved Organic Carbon to Top Consumers

Terrestrial dissolved organic carbon (DOC) provides an external carbon source to lake ecosystems. However, there is ongoing debate about whether external DOC that enters a lake can pass up the food web to support top consumers. We show, from experimental manipulation of a whole lake, that externally loaded DOC can contribute appreciably to fish biomass. Monthly additions of cane sugar with a distinct carbon stable isotope value during 2 years rapidly enriched the ¹³C content of zooplankton and macroinvertebrates, with a more gradual ¹³C enrichment of fish. After sugar addition stopped, the ¹³C content of consumers reverted towards original values. A simple isotope mixing model indicated that by the end of the sugar addition almost 20% of fish carbon in the lake was derived from the added sugar. Our results provide the first direct experimental demonstration at relevant ecological spatial and temporal scales that externally loaded DOC to lakes can indeed transfer to top consumers.     

Quantitative analysis of the importance of wind-induced circulation for the spatial structuring of planktonic populations

1. Several studies have shown that wind‐induced water movements have an important effect on the spatial distribution of crustacean zooplankton. However, few attempts have been made to quantify the effect of physical processes on these broad‐scale patterns. Much of our understanding of this spatial structure has been based on the results of isolated surveys, which do not capture the dynamic nature of the pelagic environment. 2. In this study, we have used a combination of high‐speed sampling (at a spatial resolution of 240 m) and spatial data analysis to quantify the factors influencing the horizontal spatial structure of the Daphnia galeata population in Windermere. 3. The results show that lake‐wide circulation patterns, as indicated by water temperature, account for 29–47% of the basin‐scale spatial variance in D. galeata abundance. However, these patterns are highly dynamic and change in response to the prevailing weather. This lack of temporal persistence means that the results of single‐survey sampling campaigns must be interpreted with caution.     

Ecosystem response to earlier ice break‐up date: Climate‐driven changes to water temperature,lake‐habitat‐specific production,and trout habitat and resource use

Climate warming has yielded earlier ice break‐up dates in recent decades for lakes leading to water temperature increases, altered habitat, and both increases and decreases to ecosystem productivity. Within lakes, the effect of climate warming on secondary production in littoral and pelagic habitats remains unclear. The intersection of changing habitat productivity and warming water temperatures on salmonids is important for understanding how climate warming will impact mountain ecosystems. We develop and test a conceptual model that expresses how earlier ice break‐up dates influence within lake habitat production, water temperatures and the habitat utilized by, resources obtained and behavior of salmonids in a mountain lake. We measured zoobenthic and zooplankton production from the littoral and pelagic habitats, thermal conditions, and the habitat use, resource use, and fitness of Brook Trout (Salvelinus fontinalis). We show that earlier ice break‐up conditions created a "resource‐rich" littoral–benthic habitat with increases in zoobenthic production compared to the pelagic habitat which decreased in zooplankton production. Despite the increases in littoral–benthic food resources, trout did not utilize littoral habitat or zoobenthic resources due to longer durations of warm water temperatures in the littoral zone. In addition, 87% of their resources were supported by the pelagic habitat during periods with earlier ice break‐up when pelagic resources were least abundant. The decreased reliance on littoral–benthic resources during earlier ice break‐up caused reduced fitness (mean reduction of 12 g) to trout. Our data show that changes to ice break‐up drive multi‐directional results for resource production within lake habitats and increase the duration of warmer water temperatures in food‐rich littoral habitats. The increased duration of warmer littoral water temperatures reduces the use of energetically efficient habitats culminating in decreased trout fitness.     

Water browning mediates predatory decimation of the Arctic fairy shrimp Branchinecta paludosa

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Lake browning impacts community structure and essential fatty acid content of littoral invertebrates in boreal lakes

Hydrobiologia - Many lakes in the northern hemisphere are browning due to increasing concentrations of terrestrial dissolved organic carbon (DOC). The consequences of lake browning to littoral...     

Decoupled trophic responses to long‐term recovery from acidification and associated browning in lakes

Increases in the concentration of dissolved organic matter (DOM) have been documented in many inland waters in recent decades, a process known as “browning”. Previous studies have often used space‐for‐time substitution to examine the direct consequences of increased DOM on lake ecosystems. However, browning often occurs concomitant with other ecologically important water chemistry changes that may interact with or overwhelm any potential ecological response to browning itself. Here we examine a long‐term (~20 year) dataset of 28 lakes in the Adirondack Park, New York, USA, that have undergone strong browning in response to recovery from acidification. With these data, we explored how primary producer and zooplankton consumer populations changed during this time and what physical and chemical changes best predicted these long‐term ecosystem changes. Our results indicate that changes in primary producers are likely driven by reduced water clarity due to browning, independent of changes in nutrients, counter to previously hypothesized primary producer response to browning. In contrast, declines in calcium concomitant with browning play an important role in driving long‐term declines in zooplankton biomass. Our results indicate that responses to browning at different trophic levels are decoupled from one another. Concomitant chemical changes have important implications for our understanding of the response of aquatic ecosystems to browning.     

Effects of nitrogen enrichment on zooplankton biomass and N:P recycling ratios across a DOC gradient in northern-latitude lakes

We used data from whole-lake studies to assess how changes in food quantity (phytoplankton biomass) and quality (phytoplankton community composition, seston C:P and N:P) with N fertilization affect zooplankton biomass, community composition and C:N:P stoichiometry, and their N:P recycling ratio along a gradient in lake DOC concentrations. We found that despite major differences in phytoplankton biomass with DOC (unimodal distributions, especially with N fertilization), no major differences in zooplankton biomass were detectable. Instead, phytoplankton to zooplankton biomass ratios were high, especially at intermediate DOC and after N fertilization, implying low trophic transfer efficiencies. An explanation for the observed low phytoplankton resource use, and biomass responses in zooplankton, was dominance of colony forming chlorophytes of reduced edibility at intermediate lake DOC, combined with reduced phytoplankton mineral quality (enhanced seston N:P) with N fertilization. N fertilization, however, increased zooplankton N:P recycling ratios, with largest impact at low DOC where phytoplankton benefitted from light sufficiently to cause enhanced seston N:P. Our results suggest that although N enrichment and increased phytoplankton biomass do not necessarily increase zooplankton biomass, bottom-up effects may still impact zooplankton and their N:P recycling ratio through promotion of phytoplankton species of low edibility and altered mineral quality.

     

Using functional traits to investigate the determinants of crustacean zooplankton community structure

Understanding the various processes contributing to community assembly is among the central aims of ecology. As a means of exploring this topic we quantified the relative influences of habitat filtering and competition in establishing patterns of community functional trait diversity across a landscape of lakes. Habitat filtering has been invoked in shaping community structure when co‐occurring taxa are more similar in their traits than expected by chance (under‐dispersion), and competition has been inferred as a structuring agent when co‐occurring taxa are less similar (over‐dispersion). We tested these hypotheses in crustacean zooplankton communities using a functional trait‐based approach based on five traits defining zooplankton feeding and habitat preferences across 51 lakes spanning several large limnological gradients. In general, zooplankton communities were functionally less diverse than random assemblages created from the same regional species pool. Furthermore, functional diversity was strongly correlated with variables related to lake productivity, suggesting that access to resources was the chief habitat filtering process constraining zooplankton functional diversity. This pattern was driven by the predominantly herbivorous cladocerans as opposed to the more commonly omnivorous, and sometimes carnivorous, copepods.     

Variation in spatial and temporal gradients in zooplankton spring development: the effect of climatic factors

1. We examined the temporal and spatial heterogeneity of zooplankton in lake surface waters during the spring of 3 years in Lake Washington, U.S.A., a large lake with a high production of sockeye salmon fry. 2. We show large within‐season and among‐year variation in the horizontal distribution of temperature, chlorophyll a concentration, and zooplankton in the lake. The main pattern, a delay in zooplankton population increase from the north‐ to the south‐end of the lake, recurred in each year and was persistent within each spring. 3. The delay is primarily caused by the development of a temperature gradient during spring warming, as cold mountain water enters the south end of the lake, while warm water enters the north end via a river draining a nearby lake. Climate factors, such as air temperature and precipitation during winter and spring, appear to influence the extent of the delay of zooplankton increase. 4. If the climate continues to warm, the temporal disconnection in zooplankton development between lake areas immediately influenced by cold river inflow and areas that are influenced by spring warming may increase in magnitude. Thus, the different areas of the lake may not contribute equally to fish production.     

Migrate or stay: terrestrial primary productivity and climate drive anadromy in Arctic char

A shift in the magnitude and timing of animal migrations is one of the most documented ecological effects of climate change. Although migrations are largely driven by spatial variation in resource gradients, few studies connect expected changes in primary production with geographic patterns in migratory behavior. Here, we link lake primary production to the occurrence of sea migrations in the partially anadromous salmonid Arctic char (Salvelinus alpinus L.). We compiled presence/absence records of anadromous char populations spanning productivity and temperature gradients along the Norwegian coast. The probability of anadromy decreased with increasing migration distance, maximum slope of the migration route and lake productivity. There was a significant interaction between lake productivity and migration distance. The negative effect of longer migration distances was more severe in lakes with higher productivity, indicating reduced relative profitability of migration with increased feeding opportunities in freshwater. Lake productivity was mainly driven by terrestrial primary production in the catchment. We predicted future distributions of anadromous char given downscaled temperature and precipitation changes projected by two different emission scenarios and global climate models (GCMs). Projected increases in temperature and precipitation in 2071–2100 increased terrestrial primary production and, compared to the control scenario (1961–1990), decreased the range of anadromous populations. The prevalence of anadromy decreased by 53% in the HadAm3H GCM with the A2 emission scenario, 61% in HadAm3H with the B2 scenario and 22% in ECHAM4 with the B2 scenario. Cross‐ecosystem studies (e.g., terrestrial to freshwater) are critical for understanding ecological impacts of climate change. In this case, climate‐driven increases in terrestrial primary production are expected to increase primary production in lakes and ultimately reduce the prevalence of anadromy in Arctic char populations.