Resolving the Drivers of Algal Nutrient Limitation from Boreal to Arctic Lakes and Streams

Nutrient inputs to northern freshwaters are changing, potentially altering aquatic ecosystem functioning through effects on primary producers. Yet, while primary producer growth is sensitive to nutrient supply, it is also constrained by a suite of other factors, including light and temperature, which may play varying roles across stream and lake habitats. Here, we use bioassay results from 89 lakes and streams spanning northern boreal to Arctic Sweden to test for differences in nutrient limitation status of algal biomass along gradients in colored dissolved organic carbon (DOC), water temperature, and nutrient concentrations, and to ask whether there are distinct patterns and drivers between habitats. Single nitrogen (N) limitation or primary N-limitation with secondary phosphorus (P) limitation of algal biomass was the most common condition for streams and lakes. Average response to N-addition was a doubling in biomass; however, the degree of limitation was modulated by the distinct physical and chemical conditions in lakes versus streams and across boreal to Arctic regions. Overall, algal responses to N-addition were strongest at sites with low background concentrations of dissolved inorganic N. Low temperatures constrained biomass responses to added nutrients in lakes but had weaker effects on responses in streams. Further, DOC mediated the response of algal biomass to nutrient addition differently among lakes and streams. Stream responses were dampened at higher DOC, whereas lake responses to nutrient addition increased from low to moderate DOC but were depressed at high DOC. Our results suggest that future changes in nutrient availability, particularly N, will exert strong effects on the trophic state of northern freshwaters. However, we highlight important differences in the physical and chemical factors that shape algal responses to nutrient availability in different parts of aquatic networks, which will ultimately affect the integrated response of northern aquatic systems to ongoing environmental changes

Stream metabolism controls diel patterns and evasion of CO2 in Arctic streams

Streams play an important role in the global carbon (C) cycle, accounting for a large portion of CO2 evaded from inland waters despite their small areal coverage. However, the relative importance of different terrestrial and aquatic processes driving CO2 production and evasion from streams remains poorly understood. In this study, we measured O-2 and CO2 continuously in streams draining tundra-dominated catchments in northern Sweden, during the summers of 2015 and 2016. From this, we estimated daily metabolic rates and CO2 evasion simultaneously and thus provide insight into the role of stream metabolism as a driver of C dynamics in Arctic streams. Our results show that aquatic biological processes regulate CO2 concentrations and evasion at multiple timescales. Photosynthesis caused CO2 concentrations to decrease by as much as 900 ppm during the day, with the magnitude of this diel variation being strongest at the low-turbulence streams. Diel patterns in CO2 concentrations in turn influenced evasion, with up to 45% higher rates at night. Throughout the summer, CO2 evasion was sustained by aquatic ecosystem respiration, which was one order of magnitude higher than gross primary production. Furthermore, in most cases, the contribution of stream respiration exceeded CO2 evasion, suggesting that some stream reaches serve as net sources of CO2, thus creating longitudinal heterogeneity in C production and loss within this stream network. Overall, our results provide the first link between stream metabolism and CO2 evasion in the Arctic and demonstrate that stream metabolic processes are key drivers of the transformation and fate of terrestrial organic matter exported from these landscapes.Originally included in thesis in manuscript form.</p

Persistent nitrogen limitation of stream biofilm communities along climate gradients in the Arctic

Climate change is rapidly reshaping Arctic landscapes through shifts in vegetation cover and productivity, soil resource mobilization, and hydrological regimes. The implications of these changes for stream ecosystems and food webs is unclear and will depend largely on microbial biofilm responses to concurrent shifts in temperature, light, and resource supply from land. To study those responses, we used nutrient diffusing substrates to manipulate resource supply to biofilm communities along regional gradients in stream temperature, riparian shading, and dissolved organic carbon (DOC) loading in Arctic Sweden. We found strong nitrogen (N) limitation across this gradient for gross primary production, community respiration and chlorophyll-a accumulation. For unamended biofilms, activity and biomass accrual were not closely related to any single physical or chemical driver across this region. However, the magnitude of biofilm response to N addition was: in tundra streams, biofilm response was constrained by thermal regimes, whereas variation in light availability regulated this response in birch and coniferous forest streams. Furthermore, heterotrophic responses to experimental N addition increased across the region with greater stream water concentrations of DOC relative to inorganic N. Thus, future shifts in resource supply to these ecosystems are likely to interact with other concurrent environmental changes to regulate stream productivity. Indeed, our results suggest that in the absence of increased nutrient inputs, Arctic streams will be less sensitive to future changes in other habitat variables such as temperature and DOC loading

Resolving the Drivers of Algal Nutrient Limitation from Boreal to Arctic Lakes and Streams

Nutrient inputs to northern freshwaters are changing, potentially altering aquatic ecosystem functioning through effects on primary producers. Yet, while primary producer growth is sensitive to nutrient supply, it is also constrained by a suite of other factors, including light and temperature, which may play varying roles across stream and lake habitats. Here, we use bioassay results from 89 lakes and streams spanning northern boreal to Arctic Sweden to test for differences in nutrient limitation status of algal biomass along gradients in colored dissolved organic carbon (DOC), water temperature, and nutrient concentrations, and to ask whether there are distinct patterns and drivers between habitats. Single nitrogen (N) limitation or primary N-limitation with secondary phosphorus (P) limitation of algal biomass was the most common condition for streams and lakes. Average response to N-addition was a doubling in biomass; however, the degree of limitation was modulated by the distinct physical and chemical conditions in lakes versus streams and across boreal to Arctic regions. Overall, algal responses to N-addition were strongest at sites with low background concentrations of dissolved inorganic N. Low temperatures constrained biomass responses to added nutrients in lakes but had weaker effects on responses in streams. Further, DOC mediated the response of algal biomass to nutrient addition differently among lakes and streams. Stream responses were dampened at higher DOC, whereas lake responses to nutrient addition increased from low to moderate DOC but were depressed at high DOC. Our results suggest that future changes in nutrient availability, particularly N, will exert strong effects on the trophic state of northern freshwaters. However, we highlight important differences in the physical and chemical factors that shape algal responses to nutrient availability in different parts of aquatic networks, which will ultimately affect the integrated response of northern aquatic systems to ongoing environmental changes

Resolving the Drivers of Algal Nutrient Limitation from Boreal to Arctic Lakes and Streams

Nutrient inputs to northern freshwaters are changing, potentially altering aquatic ecosystem functioning through effects on primary producers. Yet, while primary producer growth is sensitive to nutrient supply, it is also constrained by a suite of other factors, including light and temperature, which may play varying roles across stream and lake habitats. Here, we use bioassay results from 89 lakes and streams spanning northern boreal to Arctic Sweden to test for differences in nutrient limitation status of algal biomass along gradients in colored dissolved organic carbon (DOC), water temperature, and nutrient concentrations, and to ask whether there are distinct patterns and drivers between habitats. Single nitrogen (N) limitation or primary N-limitation with secondary phosphorus (P) limitation of algal biomass was the most common condition for streams and lakes. Average response to N-addition was a doubling in biomass; however, the degree of limitation was modulated by the distinct physical and chemical conditions in lakes versus streams and across boreal to Arctic regions. Overall, algal responses to N-addition were strongest at sites with low background concentrations of dissolved inorganic N. Low temperatures constrained biomass responses to added nutrients in lakes but had weaker effects on responses in streams. Further, DOC mediated the response of algal biomass to nutrient addition differently among lakes and streams. Stream responses were dampened at higher DOC, whereas lake responses to nutrient addition increased from low to moderate DOC but were depressed at high DOC. Our results suggest that future changes in nutrient availability, particularly N, will exert strong effects on the trophic state of northern freshwaters. However, we highlight important differences in the physical and chemical factors that shape algal responses to nutrient availability in different parts of aquatic networks, which will ultimately affect the integrated response of northern aquatic systems to ongoing environmental changes

Nitrogen supply and physical disturbance shapes Arctic stream nitrogen uptake through effects on metabolic activity

Climate change in the Arctic is altering the delivery of nutrients from terrestrial to aquatic ecosystems. The impact of these changes on downstream lakes and rivers is influenced by the capacity of small streams to retain such inputs. Given the potential for nutrient limitation in oligotrophic Arctic streams, biotic demand should be high, unless harsh environmental conditions maintain low biomass standing stocks that limit nutrient uptake capacity. We assessed the drivers of nutrient uptake in two contrasting headwater environments in Arctic Sweden: one stream draining upland tundra and the other draining an alluvial valley with birch forest. At both sites, we measured nitrate (NO3−) uptake biweekly using short-term slug releases and estimated rates of gross primary production (GPP) and ecosystem respiration from continuous dissolved oxygen measurements. Catchment characteristics were associated with distinct stream chemical and biological properties. For example, the tundra stream maintained relatively low NO3− concentrations (average: 46 µg N/L) and rates of GPP (0.2 g O2 m−2 day−1). By comparison, the birch forest stream was more NO3− rich (88 µg N/L) and productive (GPP: 1.7 g O2 m−2 day−1). These differences corresponded to greater areal NO3− uptake rate and increased NO3− use efficiency (as uptake velocity) in the birch forest stream (max 192 µg N m−2 min−1 and 96 mm/hr) compared to its tundra counterpart (max 52 µg N m−2 min−1 and 49 mm/hr) during 2017. Further, different sets of environmental drivers predicted temporal patterns of nutrient uptake at these sites: abiotic factors (e.g. NO3− concentration and discharge) were associated with changes in uptake in the tundra stream, while metabolic activity was more important in the birch forest stream. Between sites, variation in uptake metrics suggests that the ability to retain pulses of nutrients is linked to nutrient supply regimes controlled at larger spatial and temporal scales and habitat properties that promote biomass accrual and thus biotic demand. Overall, constraints on biotic potential imposed by the habitat template determined the capacity of these high latitude streams to respond to future changes in nutrient inputs arising from climate warming or human land use.Originally included in thesis in manuscript form.</p

Warming stimulates sediment denitrification at the expense of anaerobic ammonium oxidation.

Temperature is one of the fundamental environmental variables governing microbially mediated denitrification and anaerobic ammonium oxidation (anammox) in sediments. The GHG nitrous oxide (N2O) is produced during denitrification, but not by anammox, and knowledge of how these pathways respond to global warming remains limited. Here, we show that warming directly stimulates denitrification-derived N2O production and that the warming response for N2O production is slightly higher than the response for denitrification in subtropical sediments. Moreover, denitrification had a higher optimal temperature than anammox. Integrating our data into a global compilation indicates that denitrifiers are more thermotolerant, whereas anammox bacteria are relatively psychrotolerant. Crucially, recent summer temperatures in low-latitude sediments have exceeded the optimal temperature of anammox, implying that further warming may suppress anammox and direct more of the nitrogen flow towards denitrification and associated N2O production, leading to a positive climate feedback at low latitudes