CO2 exchange in three Canadian High Arctic ecosystems: response to long-term experimental warming

Carbon dioxide exchange, soil C and N, leaf mineral nutrition and leaf carbon isotope discrimination (LCID‐Δ) were measured in three High Arctic tundra ecosystems over 2 years under ambient and long‐term (9 years) warmed (～2°C) conditions. These ecosystems are located at Alexandra Fiord (79°N) on Ellesmere Island, Nunavut, and span a soil water gradient; dry, mesic, and wet tundra. Growing season CO₂ fluxes (i.e., net ecosystem exchange (NEE), gross ecosystem photosynthesis (GEP), and ecosystem respiration (R_e)) were measured using an infrared gas analyzer and winter C losses were estimated by chemical absorption. All three tundra ecosystems lost CO₂ to the atmosphere during the winter, ranging from 7 to 12 g CO₂‐C m^?2 season^?1 being highest in the wet tundra. The period during the growing season when mesic tundra switch from being a CO₂ source to a CO₂ sink was increased by 2 weeks because of warming and increases in GEP. Warming during the summer stimulated dry tundra GEP more than R_e and thus, NEE was consistently greater under warmed as opposed to ambient temperatures. In mesic tundra, warming stimulated GEP with no effect on R_e increasing NEE by ～10%, especially in the first half of the summer. During the ～70 days growing season (mid‐June–mid‐August), the dry and wet tundra ecosystems were net CO₂‐C sinks (30 and 67 g C m^?2 season^?1, respectively) and the mesic ecosystem was a net C source (58 g C m^?2 season^?1) to the atmosphere under ambient temperature conditions, due in part to unusual glacier melt water flooding that occurred in the mesic tundra. Experimental warming during the growing season increased net C uptake by ～12% in dry tundra, but reduced net C uptake by ～20% in wet tundra primarily because of greater rates of R_e as opposed to lower rates of GEP. Mesic tundra responded to long‐term warming with ～30% increase in GEP with almost no change in R_e reducing this tundra type to a slight C source (17 g C m^?2 season^?1). Warming caused LCID of Dryas integrafolia plants to be higher in dry tundra and lower in Salix arctic plants in mesic and wet tundra. Our findings indicate that: (1) High Arctic ecosystems, which occur in similar mesoclimates, have different net CO₂ exchange rates with the atmosphere; (2) long‐term warming can increase the net CO₂ exchange of High Arctic tundra by stimulating GEP, but it can also reduce net CO₂ exchange in some tundra types during the summer by stimulating R_e to a greater degree than stimulating GEP; (3) after 9 years of experimental warming, increases in soil carbon and nitrogen are detectable, in part, because of increases in deciduous shrub cover, biomass, and leaf litter inputs; (4) dry tundra increases in GEP, in response to long‐term warming, is reflected in D. integrifolia LCID; and (5) the differential carbon exchange responses of dry, mesic, and wet tundra to similar warming magnitudes appear to depend, in part, on the hydrologic (soil water) conditions. Annual net ecosystem CO₂‐C exchange rates ranged from losses of 64 g C m^?2 yr^?1 to gains of 55 g C m^?2 yr^?1. These magnitudes of positive NEE are close to the estimates of NPP for these tundra types in Alexandra Fiord and in other High Arctic locations based on destructive harvests.     

Greater Abundance of <Emphasis Type="Italic">Betula nana</Emphasis> and Early Onset of the Growing Season Increase Ecosystem CO<Subscript>2</Subscript> Uptake in West Greenland

Expansion of deciduous shrubs is a common observation throughout the Arctic, with implications for carbon (C) cycling. Shrubs may increase net ecosystem C uptake through greater leaf area and gross ecosystem photosynthesis (GEP), and/or through cooler summer soils and reduced ecosystem respiration (ER). We used a space-for-time substitution combined with experimental warming at a Low Arctic site in West Greenland to examine the biophysical effects of increased temperature and Betula nana abundance on ecosystem CO₂ exchange. Communities dominated by Betula were much stronger C sinks than graminoid communities due to greater GEP and lower ER. The warming treatment had little effect on GEP, ER, or net ecosystem CO₂ exchange (NEE). The start of the growing season has been advancing at our study site, as indicated by long-term observations of plant phenology. In a retrospective analysis, we estimate that earlier onset of the growing season has increased the strength of the ecosystem C sink at rates of 1.3 and 2.1 g C m^?2 y^?1 in Betula and graminoid tundra, respectively, since 2002. However, earlier, and presumably longer, growing seasons may be associated with greater potential for drought stress. Our data suggest that mid-summer drought-induced GEP declines may partially offset C gains associated with an earlier start to the growing season. Our results suggest greater deciduous shrub abundance and longer growing seasons will likely lead to greater net C uptake in our study area, while highlighting important complexities associated with drought and plant community composition.     

The interaction of biotic and abiotic factors at multiple spatial scales affects the variability of CO<Subscript>2</Subscript> fluxes in polar environments

Climate change may turn Arctic biomes from carbon sinks into sources and vice versa, depending on the balance between gross ecosystem photosynthesis, ecosystem respiration (ER) and the resulting net ecosystem exchange (NEE). Photosynthetic capacity is species specific, and thus, it is important to quantify the contribution of different target plant species to NEE and ER. At Ny Ålesund (Svalbard archipelago, Norway), we selected different Arctic tundra plant species and measured CO₂ fluxes at plot scale and photosynthetic capacity at leaf scale. We aimed to analyze trends in CO₂ fluxes during the transition seasons (beginning vs. end of the growing season) and assess which abiotic (soil temperature, soil moisture, PAR) and biotic (plot type, phenology, LAI, photosynthetic capacity) factors influenced CO₂ emissions. NEE and ER differed between vegetation communities. All communities acted as CO₂ sources, with higher source strength at the beginning than at the end of the growing season. The key factors affecting NEE were soil temperature, LAI and species-specific photosynthetic capacities, coupled with phenology. ER was always influenced by soil temperature. Measurements of photosynthetic capacity indicated different responses among species to light intensity, as well as suggesting possible gains in response to future increases in atmospheric CO₂ concentrations. Species-specific adaptation to low temperatures could trigger significant feedbacks in a climate change context. Our data highlight the need to quantify the role of dominant species in the C cycle (sinks or sources), as changes of vegetation composition or species phenology in response to climate change may have great impact on the regional CO₂ balance.     

Winter precipitation and snow accumulation drive the methane sink or source strength of Arctic tussock tundra

Arctic winter precipitation is projected to increase with global warming, but some areas will experience decreases in snow accumulation. Although Arctic CH₄ emissions may represent a significant climate forcing feedback, long‐term impacts of changes in snow accumulation on CH₄ fluxes remain uncertain. We measured ecosystem CH₄ fluxes and soil CH₄ and CO₂ concentrations and ¹³C composition to investigate the metabolic pathways and transport mechanisms driving moist acidic tundra CH₄ flux over the growing season (Jun–Aug) after 18 years of experimental snow depth increases and decreases. Deeper snow increased soil wetness and warming, reducing soil %O₂ levels and increasing thaw depth. Soil moisture, through changes in soil %O₂ saturation, determined predominance of methanotrophy or methanogenesis, with soil temperature regulating the ecosystem CH₄ sink or source strength. Reduced snow (RS) increased the fraction of oxidized CH₄ (Fox) by 75–120% compared to Ambient, switching the system from a small source to a net CH₄ sink (21 ± 2 and ?31 ± 1 mg CH₄ m^?2 season^?1 at Ambient and RS). Deeper snow reduced Fox by 35–40% and 90–100% in medium‐ (MS) and high‐ (HS) snow additions relative to Ambient, contributing to increasing the CH₄ source strength of moist acidic tundra (464 ± 15 and 3561 ± 97 mg CH₄ m^?2 season^?1 at MS and HS). Decreases in Fox with deeper snow were partly due to increases in plant‐mediated CH₄ transport associated with the expansion of tall graminoids. Deeper snow enhanced CH₄ production within newly thawed soils, responding mainly to soil warming rather than to increases in acetate fermentation expected from thaw‐induced increases in SOC availability. Our results suggest that increased winter precipitation will increase the CH₄ source strength of Arctic tundra, but the resulting positive feedback on climate change will depend on the balance between areas with more or less snow accumulation than they are currently facing.     

Analyzing the Effects of Growing Season Length on the Net Ecosystem Production of an Alpine Grassland Using Model–Data Fusion

Alpine ecosystems are, similar to arctic ecosystems, characterized by a very long snow season. Previous studies investigating arctic or alpine ecosystems have shown that winter CO₂ effluxes can dominate the annual balance and that the timing and duration of the snow cover plays a crucial role for plant growth and phenology and might also influence the growing season ecosystem CO₂ strength and dynamics. The objective of this study was to analyze seasonal and annual CO₂ balances of a grassland site at an elevation of 2440 m a.s.l in the Swiss central Alps. We continuously measured the NEP using the eddy covariance method from June 2013 to October 2014, covering two growing seasons and one winter. We analyzed the influence of snow melt date on the CO₂ exchange dynamics at this site, because snow melt differed about 24 days between the 2 years. To this end, we employed a process-based ecosystem carbon cycling model to disentangle the co-occurring effects of growing season length, environmental conditions during the growing season, and physiological/structural properties of the canopy on the ecosystem carbon balance. During the measurement period, the site was a net sink for CO₂ although winter efflux contributed significantly to the total balance. The cumulative growing season NEP as well as mean and maximum daily CO₂ uptake rates was lower during the year with the later snow melt, and the results indicated that the differences were mainly due to differing growing season lengths.     

Microbial Respiration in Arctic Upland and Peat Soils as a Source of Atmospheric Carbon Dioxide

Knowledge on soil microbial respiration (SMR) rates and thus soil-related CO₂ losses from Arctic soils is vital because of the crucial importance of this ecosystem within the global carbon (C) cycle and climate system. Here, we measured SMR from various habitats during the growing season in Russian subarctic tundra by applying two different approaches: ¹⁴C partitioning approach and root trenching. The variable habitats encompassed peat and mineral soils, bare and vegetated surfaces and included both dry and moist ones. The field experiment was complemented by laboratory studies to measure bioavailability of soil carbon and identify sources of CO₂. Differences in bioavailability of soils, measured in the laboratory as basal soil respiration rates, were generally greater than inter-site differences in SMR rates measured in situ, suggesting secondary constraints at field conditions, such as soil C content. There was a tendency towards lower SMR in vegetated peat plateaus compared to upland mineral tundra (on average 137 vs. 185 g CO₂ m^?2 growing season^?1, respectively), but no significant differences were found. Surprisingly, the bare surfaces (peat circles) with 3500-year-old C at the surface exhibited about the largest SMR among all sites as shown by both methods. This was related to the general development of peat plateaus in the region, and uplifting of deeper peat with high C content to the surface during the genesis of peat circles. This observation is particularly relevant for decomposition of deeper peat in vegetated peat plateaus, where soil material similar to the bare surfaces can be found. The data indicate that the large stocks of C stored in permafrost peatlands are principally available for decomposition despite old age.     

The effects of water table manipulation and elevated temperature on the net CO2 flux of wet sedge tundra ecosystems

In situ manipulations were conducted in a naturally drained lake on the arctic coastal plain near Prudhoe Bay, Alaska (70 °21.98′ N, 148 °33.72′ W) to assess the potential short-term effects of decreased water table and elevated temperature on net ecosystem CO₂ flux. The experiments were conducted over a 2-year period, and during that time, water table depth of drained plots was maintained on average 7 cm lower than the ambient water table, and surface temperatures of plots exposed to elevated temperature were increased on average 0.5 °C. Water table drainage, and to a lesser extent elevated temperature, resulted in significant increases in ecosystem respiration (ER) rates, and only small and variable changes in gross ecosystem productivity (GEP). As a result, drained plots were net sources of ≈ 40 gC m^–2 season^–1 over both years of manipulation, while control plots were net sinks of atmospheric CO₂ of about 10 gC m^–2 season^–1 (growing season length was an estimated 125 days). Control plots exposed to elevated temperatures accumulated slightly more carbon than control plots exposed to ambient temperatures. The direct effects of elevated temperature on net CO₂ flux, ER, and GEP were small, however, elevated temperature appeared to interact with drainage to exacerbate the amount of net carbon loss. These data suggest that many currently saturated or nearly saturated wet sedge ecosystems of the north slope of Alaska may become significant sources of CO₂ to the atmosphere if climate change predictions of increased evapotranspiration and reduced soil water status are realized. There is ample evidence that this may be already occurring in arctic Alaska, as a change in net carbon balance has been observed for both tussock and wet-sedge tundra ecosystems over the last 2–3 decades, which coincides with a recent increase in surface temperature and an associated decrease in soil water content. In contrast, if precipitation increases relatively more than evapotranspiration, then increases in soil moisture content will likely result in greater carbon accumulation.     

Stimulation of both photosynthesis and respiration in response to warmer and drier conditions in a boreal peatland ecosystem

Peatland ecosystems have been consistent carbon (C) sinks for millennia, but it has been predicted that exposure to warmer temperatures and drier conditions associated with climate change will shift the balance between ecosystem photosynthesis and respiration providing a positive feedback to atmospheric CO₂ concentration. Our main objective was to determine the sensitivity of ecosystem photosynthesis, respiration and net ecosystem production (NEP) measured by eddy covariance, to variation in temperature and water table depth associated with interannual shifts in weather during 2004–2009. Our study was conducted in a moderately rich treed fen, the most abundant peatland type in western Canada, in a region (northern Alberta) where peatland ecosystems are a significant landscape component. During the study, the average growing season (May–October) water depth declined approximately 38 cm, and temperature [expressed as cumulative growing degree days (GDD, March–October)] varied approximately 370 GDD. Contrary to previous predictions, both ecosystem photosynthesis and respiration showed similar increases in response to warmer and drier conditions. The ecosystem remained a strong net sink for CO₂ with an average NEP (± SD) of 189 ± 47 g C m^?2 yr^?1. The current net CO₂ uptake rates were much higher than C accumulation in peat determined from analyses of the relationship between peat age and cumulative C stock. The balance between C addition to, and total loss from, the top 0–30 cm depth (peat age range 0–70 years) of shallow peat cores averaged 43 ± 12 g C m^?2 yr^?1. The apparent long‐term average rate of net C accumulation in basal peat samples was 19–24 g C m^?2 yr^?1. The difference between current rates of net C uptake and historical rates of peat accumulation is likely a result of vegetation succession and recent increases in tree establishment and productivity.     

Soil–plant N processes in a High Arctic ecosystem,NW Greenland are altered by long‐term experimental warming and higher rainfall

Rapid temperature and precipitation changes in High Arctic tundra ecosystems are altering the biogeochemical cycles of carbon (C) and nitrogen (N), but in ways that are difficult to predict. The challenge grows from the uncertainty of N cycle responses and the extent to which shifts in soil N are coupled with the C cycle and productivity of tundra systems. We used a long‐term (since 2003) experiment of summer warming and supplemental summer water additions to a High Arctic ecosystem in NW Greenland, and applied a combination of discrete sampling and in situ soil core incubations to measure C and N pools and seasonal microbial processes that might control plant‐available N. We hypothesized that elevated temperature and increased precipitation would stimulate microbial activity and net inorganic N mineralization, thereby increasing plant N‐availability through the growing season. While we did find increased N mineralization rates under both global change scenarios, water addition also significantly increased net nitrification rates, loss of NO₃^?‐N via leaching, and lowered rates of labile organic N production. We also expected the chronic warming and watering would lead to long‐term changes in soil N‐cycling that would be reflected in soil δ¹⁵N values. We found that soil δ¹⁵N decreased under the different climate change scenarios. Our results suggest that temperature accelerates biological processes and existing C and N transformations, but moisture increases soil hydraulic connectivity and so alters the pathways, and changes the fate of the products of C and N transformations. In addition, our findings indicate that warmer, wetter High Arctic tundra will be cycling N and C in ways that may transform these landscapes in part leading to greater C sequestration, but simultaneously, N losses from the upper soil profile that may be transported to depth dissolved in water and or transported off site in lateral flow.     

Land‐atmosphere exchange of methane from soil thawing to soil freezing in a high‐Arctic wet tundra ecosystem

The land‐atmosphere exchange of methane (CH₄) and carbon dioxide (CO₂) in a high‐Arctic wet tundra ecosystem (Rylekærene) in Zackenberg, north‐eastern Greenland, was studied over the full growing season and until early winter in 2008 and from before snow melt until early winter in 2009. The eddy covariance technique was used to estimate CO₂ fluxes and a combination of the gradient and eddy covariance methods was used to estimate CH₄ fluxes. Small CH₄ bursts were observed during spring thawing 2009, but these existed during short periods and would not have any significant effect on the annual budget. Growing season CH₄ fluxes were well correlated with soil temperature, gross primary production, and active layer thickness. The CH₄ fluxes remained low during the entire autumn, and until early winter. No increase in CH₄ fluxes were seen as the soil started to freeze. However, in autumn 2008 there were two CH₄ burst events that were highly correlated with atmospheric turbulence. They were likely associated with the release of stored CH₄ from soil and vegetation cavities. Over the measurement period, 7.6 and 6.5 g C m^?2 was emitted as CH₄ in 2008 and in 2009, respectively. Rylekærene acted as a C source during the warmer and wetter measurement period 2008, whereas it was a C sink for the colder and drier period of 2009. Wet tundra ecosystems, such as Rylekærene may thus play a more significant role for the climate in the future, as temperature and precipitation are predicted to increase in the high‐Arctic.     

Effects of experimental warming of air,soil and permafrost on carbon balance in Alaskan tundra

The carbon (C) storage capacity of northern latitude ecosystems may diminish as warming air temperatures increase permafrost thaw and stimulate decomposition of previously frozen soil organic C. However, warming may also enhance plant growth so that photosynthetic carbon dioxide (CO₂) uptake may, in part, offset respiratory losses. To determine the effects of air and soil warming on CO₂ exchange in tundra, we established an ecosystem warming experiment – the Carbon in Permafrost Experimental Heating Research (CiPEHR) project – in the northern foothills of the Alaska Range in Interior Alaska. We used snow fences coupled with spring snow removal to increase deep soil temperatures and thaw depth (winter warming) and open‐top chambers to increase growing season air temperatures (summer warming). Winter warming increased soil temperature (integrated 5–40 cm depth) by 1.5 °C, which resulted in a 10% increase in growing season thaw depth. Surprisingly, the additional 2 kg of thawed soil C m^?2 in the winter warming plots did not result in significant changes in cumulative growing season respiration, which may have been inhibited by soil saturation at the base of the active layer. In contrast to the limited effects on growing‐season C dynamics, winter warming caused drastic changes in winter respiration and altered the annual C balance of this ecosystem by doubling the net loss of CO₂ to the atmosphere. While most changes to the abiotic environment at CiPEHR were driven by winter warming, summer warming effects on plant and soil processes resulted in 20% increases in both gross primary productivity and growing season ecosystem respiration and significantly altered the age and sources of CO₂ respired from this ecosystem. These results demonstrate the vulnerability of organic C stored in near surface permafrost to increasing temperatures and the strong potential for warming tundra to serve as a positive feedback to global climate change.     

Short-term impacts of active layer detachments on carbon exchange in a High Arctic ecosystem,Cape Bounty,Nunavut, Canada

Localized permafrost disturbances such as active layer detachments (ALDs) are increasing in frequency and severity across the Canadian Arctic impacting terrestrial ecosystem functioning. However, the contribution of permafrost disturbance-carbon feedbacks to the carbon (C) balance of Arctic ecosystems is poorly understood. Here, we explore the short-term impact of active layer detachments (ALDs) on carbon dioxide (CO₂) exchange in a High Arctic semi-desert ecosystem by comparing midday C exchange between undisturbed areas, moderately disturbed areas (intact islands of vegetation within an ALD), and highly disturbed areas (non-vegetated areas due to ALD). Midday C exchange was measured using a static chamber method between June 23 and August 8 during the 2009 and 2010 growing seasons. Results show that areas of high disturbance had significantly reduced gross ecosystem exchange and ecosystem respiration (R _E) compared to control and moderately disturbed areas. Moderately disturbed areas showed significantly enhanced net ecosystem exchange compared to areas of high disturbance, but were not significantly different from control areas. Disturbance did not significantly impact soil thermal, physical or chemical properties. According to average midday fluxes, ALDs as a whole (moderately disturbed areas: ?1.942 μmol m^?2 s^?1+ highly disturbed areas: 2.969 μmol m^?2 s^?1) were a small CO₂ source of 1.027 μmol m^?2 s^?1 which did not differ significantly from average midday fluxes in control areas 1.219 μmol m^?2 s^?1. The findings of this study provide evidence that the short-term impacts of ALDs on midday, net C exchange and soil properties in a High Arctic semi-desert are minimal.     

Long-Term Release of Carbon Dioxide from Arctic Tundra Ecosystems in Alaska

Releases of the greenhouse gases carbon dioxide (CO₂) and methane (CH₄) from thawing permafrost are expected to be among the largest feedbacks to climate from arctic ecosystems. However, the current net carbon (C) balance of terrestrial arctic ecosystems is unknown. Recent studies suggest that these ecosystems are sources, sinks, or approximately in balance at present. This uncertainty arises because there are few long-term continuous measurements of arctic tundra CO₂ fluxes over the full annual cycle. Here, we describe a pattern of CO₂ loss based on the longest continuous record of direct measurements of CO₂ fluxes in the Alaskan Arctic, from two representative tundra ecosystems, wet sedge and heath tundra. We also report on a shorter time series of continuous measurements from a third ecosystem, tussock tundra. The amount of CO₂ loss from both heath and wet sedge ecosystems was related to the timing of freeze-up of the soil active layer in the fall. Wet sedge tundra lost the most CO₂ during the anomalously warm autumn periods of September–December 2013–2015, with CH₄ emissions contributing little to the overall C budget. Losses of C translated to approximately 4.1 and 1.4% of the total soil C stocks in active layer of the wet sedge and heath tundra, respectively, from 2008 to 2015. Increases in air temperature and soil temperatures at all depths may trigger a new trajectory of CO₂ release, which will be a significant feedback to further warming if it is representative of larger areas of the Arctic.     

Ecosystem nitrogen fixation throughout the snow‐free period in subarctic tundra: effects of willow and birch litter addition and warming

Nitrogen (N) fixation in moss‐associated cyanobacteria is one of the main sources of available N for N‐limited ecosystems such as subarctic tundra. Yet, N₂ fixation in mosses is strongly influenced by soil moisture and temperature. Thus, temporal scaling up of low‐frequency in situ measurements to several weeks, months or even the entire growing season without taking into account changes in abiotic conditions cannot capture the variation in moss‐associated N₂ fixation. We therefore aimed to estimate moss‐associated N₂ fixation throughout the snow‐free period in subarctic tundra in field experiments simulating climate change: willow (Salix myrsinifolia) and birch (Betula pubescens spp. tortuosa) litter addition, and warming. To achieve this, we established relationships between measured in situ N₂ fixation rates and soil moisture and soil temperature and used high‐resolution measurements of soil moisture and soil temperature (hourly from May to October) to model N₂ fixation. The modelled N₂ fixation rates were highest in the warmed (2.8 ± 0.3 kg N ha^?1) and birch litter addition plots (2.8 ± 0.2 kg N ha^?1), and lowest in the plots receiving willow litter (1.6 ± 0.2 kg N ha^?1). The control plots had intermediate rates (2.2 ± 0.2 kg N ha^?1). Further, N₂ fixation was highest during the summer in the warmed plots, but was lowest in the litter addition plots during the same period. The temperature and moisture dependence of N₂ fixation was different between the climate change treatments, indicating a shift in the N₂ fixer community. Our findings, using a combined empirical and modelling approach, suggest that a longer snow‐free period and increased temperatures in a future climate will likely lead to higher N₂ fixation rates in mosses. Yet, the consequences of increased litter fall on moss‐associated N₂ fixation due to shrub expansion in the Arctic will depend on the shrub species’ litter traits.     

Performance of High Arctic tundra plants improved during but deteriorated after exposure to a simulated extreme temperature event

Arctic ecosystems are known to be extremely vulnerable to climate change. As the Intergovernmental Panel on Climate Change scenarios project extreme climate events to increase in frequency and severity, we exposed High Arctic tundra plots during 8 days in summer to a temperature rise of approximately 9°C, induced by infrared irradiation, followed by a recovery period. Increased plant growth rates during the heat wave, increased green cover at the end of the heat wave and higher chlorophyll concentrations of all four predominating species (Salix arctica Pall., Arctagrostis latifolia Griseb., Carex bigelowii Torr. ex Schwein and Polygonum viviparum L.) after the recovery period, indicated stimulation of vegetative growth. Improved plant performance during the heat wave was confirmed at plant level by higher leaf photochemical efficiency (F_v/F_m) and at ecosystem level by increased gross canopy photosynthesis. However, in the aftermath of the temperature extreme, the heated plants were more stressed than the unheated plants, probably because they acclimated to warmer conditions and experienced the return to (low) ambient as stressful. We also calculated the impact of the heat wave on the carbon balance of this tundra ecosystem. Below‐ and aboveground respiration were stimulated by the instantaneous warmer soil and canopy, respectively, outweighing the increased gross photosynthesis. As a result, during the heat wave, the heated plots were a smaller sink compared with their unheated counterparts, whereas afterwards the balance was not affected. If other High Arctic tundra ecosystems react similarly, more frequent extreme temperature events in a future climate may shift this biome towards a source. It is uncertain, however, whether these short‐term effects will hold when C exchange rates acclimate to higher average temperatures.     

Subalpine grassland carbon dioxide fluxes indicate substantial carbon losses under increased nitrogen deposition,but not at elevated ozone concentration

Ozone (O₃) and nitrogen (N) deposition affect plant carbon (C) dynamics and may change ecosystem C‐sink/‐source properties. We studied effects of increased background [O₃] (up to [ambient] × 2) and increased N deposition (up to +50 kg ha^?1 a^?1) on mature, subalpine grassland during the third treatment year. During 10 days and 13 nights, distributed evenly over the growth period of 2006, we measured ecosystem‐level CO₂ exchange using a static cuvette. Light dependency of gross primary production (GPP) and temperature dependency of ecosystem respiration rates (R_eco) were established. Soil temperature, soil water content, and solar radiation were monitored. Using R_eco and GPP values, we calculated seasonal net ecosystem production (NEP), based on hourly averages of global radiation and soil temperature. Differences in NEP were compared with differences in soil organic C after 5 years of treatment. The high [O₃] had no effect on aboveground dry matter productivity (DM), but seasonal mean rates of both R_eco and GPP decreased ca. 8%. NEP indicated an unaltered growing season CO₂–C balance. High N treatment, with a +31% increase in DM, mean R_eco increased ca. 3%, but GPP decreased ca. 4%. Consequently, seasonal NEP yielded a 53.9 g C m^?2 (±22.05) C loss compared with control. Independent of treatment, we observed a negative NEP of 146.4 g C m^?2 (±15.3). Carbon loss was likely due to a transient management effect, equivalent to a shift from pasture to hay meadow and a drought effect, specific to the 2006 summer climate. We argue that this resulted from strongly intensified soil microbial respiration, following mitigation of nutrient limitation. There was no interaction between O₃ and N treatments. Thus, during the 2006 growing season, the subalpine grassland lost >2% of total topsoil organic C as respired CO₂, with increased N deposition responsible for one‐third of that loss.     

Response of terrestrial carbon uptake to climate interannual variability in China

The interest in national terrestrial ecosystem carbon budgets has been increasing because the Kyoto Protocol has included some terrestrial carbon sinks in a legally binding framework for controlling greenhouse gases emissions. Accurate quantification of the terrestrial carbon sink must account the interannual variations associated with climate variability and change. This study used a process‐based biogeochemical model and a remote sensing‐based production efficiency model to estimate the variations in net primary production (NPP), soil heterotrophic respiration (HR), and net ecosystem production (NEP) caused by climate variability and atmospheric CO₂ increases in China during the period 1981–2000. The results show that China's terrestrial NPP varied between 2.86 and 3.37 Gt C yr^?1 with a growth rate of 0.32% year^?1 and HR varied between 2.89 and 3.21 Gt C yr^?1 with a growth rate of 0.40% year^?1 in the period 1981–1998. Whereas the increases in HR were related mainly to warming, the increases in NPP were attributed to increases in precipitation and atmospheric CO₂. Net ecosystem production (NEP) varied between ?0.32 and 0.25 Gt C yr^?1 with a mean value of 0.07 Gt C yr^?1, leading to carbon accumulation of 0.79 Gt in vegetation and 0.43 Gt in soils during the period. To the interannual variations in NEP changes in NPP contributed more than HR in arid northern China but less in moist southern China. NEP had no a statistically significant trend, but the mean annual NEP for the 1990s was lower than for the 1980s as the increases in NEP in southern China were offset by the decreases in northern China. These estimates indicate that China's terrestrial ecosystems were taking up carbon but the capacity was undermined by the ongoing climate change. The estimated NEP related to climate variation and atmospheric CO₂ increases may account for from 40 to 80% to the total terrestrial carbon sink in China.     

Pan-Arctic soil moisture control on tundra carbon sequestration and plant productivity

Long-term atmospheric CO₂ concentration records have suggested a reduction in the positive effect of warming on high-latitude carbon uptake since the 1990s. A variety of mechanisms have been proposed to explain the reduced net carbon sink of northern ecosystems with increased air temperature, including water stress on vegetation and increased respiration over recent decades. However, the lack of consistent long-term carbon flux and in situ soil moisture data has severely limited our ability to identify the mechanisms responsible for the recent reduced carbon sink strength. In this study, we used a record of nearly 100 site-years of eddy covariance data from 11 continuous permafrost tundra sites distributed across the circumpolar Arctic to test the temperature (expressed as growing degree days, GDD) responses of gross primary production (GPP), net ecosystem exchange (NEE), and ecosystem respiration (ER) at different periods of the summer (early, peak, and late summer) including dominant tundra vegetation classes (graminoids and mosses, and shrubs). We further tested GPP, NEE, and ER relationships with soil moisture and vapor pressure deficit to identify potential moisture limitations on plant productivity and net carbon exchange. Our results show a decrease in GPP with rising GDD during the peak summer (July) for both vegetation classes, and a significant relationship between the peak summer GPP and soil moisture after statistically controlling for GDD in a partial correlation analysis. These results suggest that tundra ecosystems might not benefit from increased temperature as much as suggested by several terrestrial biosphere models, if decreased soil moisture limits the peak summer plant productivity, reducing the ability of these ecosystems to sequester carbon during the summer.     

Carbon balance in the tundra, boreal forest and humid tropical forest during climate change: scaling up from leaf physiology and soil carbon dynamics

Carbon exchange by the terrestrial biosphere is thought to have changed since pre-industrial times in response to increasing concentrations of atmospheric CO₂ and variations (anomalies) in inter-annual air temperatures. However, the magnitude of this response, particularly that of various ecosystem types (biomes), is uncertain. Terrestrial carbon models can be used to estimate the direction and size of the terrestrial responses expected, providing that these models have a reasonable theoretical base. We formulated a general model of ecosystem carbon fluxes by linking a process-based canopy photosynthesis model to the Rothamsted soil carbon model for biomes that are not significantly affected by water limitation. The difference between net primary production (NPP) and heterotrophic soil respiration (R_h) represents net ecosystem production (NEP). The model includes (i) multiple compartments for carbon storage in vegetation and soil organic matter, (ii) the effects of seasonal changes in environmental parameters on annual NEP, and (iii) the effects of inter-annual temperature variations on annual NEP. Past, present and projected changes in atmospheric CO₂ concentration and surface air temperature (at different latitudes) were analysed for their effects on annual NEP in tundra, boreal forest and humid tropical forest biomes. In all three biomes, annual NEP was predicted to increase with CO₂ concentration but to decrease with warming. As CO₂ concentrations and temperatures rise, the positive carbon gains through increased NPP are often outweighed by losses through increased R_h, particularly at high latitudes where global warming has been (and is expected to be) most severe. We calculated that, several times during the past 140 years, both the tundra and boreal forest biomes have switched between being carbon sources (annual NEP negative) and being carbon sinks (annual NEP positive). Most recently, significant warming at high latitudes during 1988 and 1990 caused the tundra and boreal forests to be net carbon sources. Humid tropical forests generally have been a carbon sink since 1960. These modelled responses of the various biomes are in agreement with other estimates from either field measurements or geochemical models. Under projected CO₂ and temperature increases, the tundra and boreal forests will emit increasingly more carbon to the atmosphere while the humid tropical forest will continue to store carbon. Our analyses also indicate that the relative increase in the seasonal amplitude of the accumulated NEP within a year is about 0–14% year^?1 for boreal forests and 0–23% year^?1 in the tundra between 1960 and 1990.     

Historical and projected carbon balance of mature black spruce ecosystems across North America: the role of carbon–nitrogen interactions

The role of carbon (C) and nitrogen (N) interactions on sequestration of atmospheric CO₂ in black spruce ecosystems across North America was evaluated with the Terrestrial Ecosystem Model (TEM) by applying parameterizations of the model in which C–N dynamics were either coupled or uncoupled. First, the performance of the parameterizations, which were developed for the dynamics of black spruce ecosystems at the Bonanza Creek Long-Term Ecological Research site in Alaska, were evaluated by simulating C dynamics at eddy correlation tower sites in the Boreal Ecosystem Atmosphere Study (BOREAS) for black spruce ecosystems in the northern study area (northern site) and the southern study area (southern site) with local climate data. We compared simulated monthly growing season (May to September) estimates of gross primary production (GPP), total ecosystem respiration (RESP), and net ecosystem production (NEP) from 1994 to 1997 to available field-based estimates at both sites. At the northern site, monthly growing season estimates of GPP and RESP for the coupled and uncoupled simulations were highly correlated with the field-based estimates (coupled: R ²= 0.77, 0.88 for GPP and RESP; uncoupled: R ² = 0.67, 0.92 for GPP and RESP). Although the simulated seasonal pattern of NEP generally matched the field-based data, the correlations between field-based and simulated monthly growing season NEP were lower (R ² = 0.40, 0.00 for coupled and uncoupled simulations, respectively) in comparison to the correlations between field-based and simulated GPP and RESP. The annual NEP simulated by the coupled parameterization fell within the uncertainty of field-based estimates in two of three years. On the other hand, annual NEP simulated by the uncoupled parameterization only fell within the field-based uncertainty in one of three years. At the southern site, simulated NEP generally matched field-based NEP estimates, and the correlation between monthly growing season field-based and simulated NEP (R ² = 0.36, 0.20 for coupled and uncoupled simulations, respectively) was similar to the correlations at the northern site. To evaluate the role of N dynamics in C balance of black spruce ecosystems across North America, we simulated historical and projected C dynamics from 1900 to 2100 with a global-based climatology at 0.5° resolution (latitude × longitude) with both the coupled and uncoupled parameterizations of TEM. From analyses at the northern site, several consistent patterns emerge. There was greater inter-annual variability in net primary production (NPP) simulated by the uncoupled parameterization as compared to the coupled parameterization, which led to substantial differences in inter-annual variability in NEP between the parameterizations. The divergence between NPP and heterotrophic respiration was greater in the uncoupled simulation, resulting in more C sequestration during the projected period. These responses were the result of fundamentally different responses of the coupled and uncoupled parameterizations to changes in CO₂ and climate.