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.     

Effects of drainage and temperature on carbon balance of tussock tundra micrososms

We examined the importance of temperature (7°C or 15°C) and soil moisture regime (saturated or field capacity) on the carbon (C) balance of arctic tussock tundra microcosms (intact blocks of soil and vegetation) in growth chambers over an 81-day simulated growing season. We measured gaseous CO₂ exchanges, methane (CH₄) emissions, and dissolved C losses on intact blocks of tussock (Eriophorum vaginatum) and intertussock (moss-dominated). We hypothesized that under increased temperature and/or enhanced drainage, C losses from ecosystem respiration (CO₂ respired by plants and heterotrophs) would exceed gains from gross photosynthesis causing tussock tundra to become a net source of C to the atmosphere. The field capacity moisture regime caused a decrease in net CO₂ storage (NEP) in tussock tundra micrososms. This resulted from a stimulation of ecosystem respiration (probably mostly microbial) with enhanced drainage, rather than a decrease in gross photosynthesis. Elevated temperature alone had no effect on NEP because CO₂ losses from increased ecosystem respiration at elevated temperature were compensated by increased CO₂ uptake (gross photosynthesis). Although CO₂ losses from ecosystem respiration were primarily limited by drainage, CH₄ emissions, in contrast, were dependent on temperature. Furthermore, substantial dissolved C losses, especially organic C, and important microhabitat differences must be considered in estimating C balance for the tussock tundra system. As much as 20% of total C fixed in photosynthesis was lost as dissolved organic C. Tussocks stored 2x more C and emitted 5x more methane than intertussocks. In spite of the limitations of this microcosm experiment, this study has further elucidated the critical role of soil moisture regime and dissolved C losses in regulating net C balance of arctic tussock tundra.     

Individual and interactive effects of warming and nitrogen supply on CO2 fluxes and carbon allocation in subarctic grassland

Climate warming has been suggested to impact high latitude grasslands severely, potentially causing considerable carbon (C) losses from soil. Warming can also stimulate nitrogen (N) turnover, but it is largely unclear whether and how altered N availability impacts belowground C dynamics. Even less is known about the individual and interactive effects of warming and N availability on the fate of recently photosynthesized C in soil. On a 10-year geothermal warming gradient in Iceland, we studied the effects of soil warming and N addition on CO₂ fluxes and the fate of recently photosynthesized C through CO₂ flux measurements and a ¹³CO₂ pulse-labeling experiment. Under warming, ecosystem respiration exceeded maximum gross primary productivity, causing increased net CO₂ emissions. N addition treatments revealed that, surprisingly, the plants in the warmed soil were N limited, which constrained primary productivity and decreased recently assimilated C in shoots and roots. In soil, microbes were increasingly C limited under warming and increased microbial uptake of recent C. Soil respiration was increased by warming and was fueled by increased belowground inputs and turnover of recently photosynthesized C. Our findings suggest that a decade of warming seemed to have induced a N limitation in plants and a C limitation by soil microbes. This caused a decrease in net ecosystem CO₂ uptake and accelerated the respiratory release of photosynthesized C, which decreased the C sequestration potential of the grassland. Our study highlights the importance of belowground C allocation and C-N interactions in the C dynamics of subarctic ecosystems in a warmer world.     

Net ecosystem CO2 exchange and plant biomass responses to warming and N addition in a grass-dominated system during two years of net CO2 efflux

Background and Aims

Climate warming and increased atmospheric nitrogen (N) deposition both have the potential to increase plant productivity over the next century, yet they can also increase decomposition and respiration. Our aim was to examine the extent to which warming and N addition can, on balance, alter net ecosystem CO₂ exchange (NEE) in a grass-dominated system.

Methods

We measured NEE responses to warming and N addition over two growing seasons in a temperate old field using steady-state flow-through chambers, which allowed for the integrated measurement of respiration and photoassimilation effects on net CO₂ flux over diel periods. We also assessed the relationship between NEE and plant biomass responses to the warming and N treatments.

Results

In both years, our study system was a net source of carbon (C) during the snow-free season. N addition did not significantly affect diel NEE or dark respiration in either year, despite a doubling in aboveground plant biomass in response to N addition in the second year, and a corresponding increase in peak daily net CO₂ photoassimilation in N addition plots. The warming treatment also had no significant effect on NEE, although the flow-through chambers required warming to be temporarily halted during NEE measurements.

Conclusions

Overall, our results both highlight the potential divergence of plant and soil responses to N addition and demonstrate the capacity for a grass-dominated system to function as a net source of C in consecutive years.     

Influence of High Temperature on End-of-Season Tundra CO2 Exchange

The high-arctic terrestrial environment is generally recognized as one of the world's most sensitive areas with regard to global warming. In this study, we examined the influence of an isolated warm period on net ecosystem carbon dioxide (CO₂) exchange at high latitude during autumn. Using the Free Air Temperature Increase (FATI) technique, we manipulated air, soil, and vegetation temperatures in late August in a tundra site at Zackenberg in the National Park of North and East Greenland (74°N 21°W). The consequences for gross canopy photosynthesis, canopy respiration, and belowground respiration of increasing these temperatures by approximately 2.5°C were determined with closed dynamic CO₂ exchange systems. Under current temperatures, the ecosystem acted as a net CO₂ source, releasing 19 g CO₂-C m⁻² over the 14-day study period. Warm soils and senescing vegetation in autumn were unequivocally responsible for this efflux. Heating enhanced CO₂ efflux to 29 g CO₂-C m⁻². This effect was attributed to a 39% increase in belowground respiration, which was the main component of the carbon (C) budget. Gross photosynthesis, on the other hand, was not affected significantly by the simulated warming. Although the aftereffects of an isolated warm period on the C balance in early winter could be significant, simulations with a simple C budget model suggest that soil carbon pools are not affected to a great extent by such a climatic disturbance. The influence on atmospheric carbon, however, appears to be significant. Received 9 June 2000; accepted 20 December 2000.     

Summer temperature increase has distinct effects on the ectomycorrhizal fungal communities of moist tussock and dry tundra in Arctic Alaska

Arctic regions are experiencing the greatest rates of climate warming on the planet and marked changes have already been observed in terrestrial arctic ecosystems. While most studies have focused on the effects of warming on arctic vegetation and nutrient cycling, little is known about how belowground communities, such as fungi root‐associated, respond to warming. Here, we investigate how long‐term summer warming affects ectomycorrhizal (ECM) fungal communities. We used Ion Torrent sequencing of the rDNA internal transcribed spacer 2 (ITS2) region to compare ECM fungal communities in plots with and without long‐term experimental warming in both dry and moist tussock tundra. Cortinarius was the most OTU‐rich genus in the moist tundra, while the most diverse genus in the dry tundra was Tomentella. On the diversity level, in the moist tundra we found significant differences in community composition, and a sharp decrease in the richness of ECM fungi due to warming. On the functional level, our results indicate that warming induces shifts in the extramatrical properties of the communities, where the species with medium‐distance exploration type seem to be favored with potential implications for the mobilization of different nutrient pools in the soil. In the dry tundra, neither community richness nor community composition was significantly altered by warming, similar to what had been observed in ECM host plants. There was, however, a marginally significant increase in OTUs identified as ECM fungi with the medium‐distance exploration type in the warmed plots. Linking our findings of decreasing richness with previous results of increasing ECM fungal biomass suggests that certain ECM species are favored by warming and may become more abundant, while many other species may go locally extinct due to direct or indirect effects of warming. Such compositional shifts in the community might affect nutrient cycling and soil organic C storage.     

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.     

Effects of Experimental Water Table and Temperature Manipulations on Ecosystem CO2 Fluxes in an Alaskan Rich Fen

Peatlands store 30% of the world’s terrestrial soil carbon (C) and those located at northern latitudes are expected to experience rapid climate warming. We monitored growing season carbon dioxide (CO₂) fluxes across a factorial design of in situ water table (control, drought, and flooded plots) and soil warming (control vs. warming via open top chambers) treatments for 2 years in a rich fen located just outside the Bonanza Creek Experimental Forest in interior Alaska. The drought (lowered water table position) treatment was a weak sink or small source of atmospheric CO₂ compared to the moderate atmospheric CO₂ sink at our control. This change in net ecosystem exchange was due to lower gross primary production and light-saturated photosynthesis rather than increased ecosystem respiration. The flooded (raised water table position) treatment was a greater CO₂ sink in 2006 due largely to increased early season gross primary production and higher light-saturated photosynthesis. Although flooding did not have substantial effects on rates of ecosystem respiration, this water table treatment had lower maximum respiration rates and a higher temperature sensitivity of ecosystem respiration than the control plot. Surface soil warming increased both ecosystem respiration and gross primary production by approximately 16% compared to control (ambient temperature) plots, with no net effect on net ecosystem exchange. Results from this rich fen manipulation suggest that fast responses to drought will include reduced ecosystem C storage driven by plant stress, whereas inundation will increase ecosystem C storage by stimulating plant growth.     

Nonlinear CO2 flux response to 7 years of experimentally induced permafrost thaw

Rapid Arctic warming is expected to increase global greenhouse gas concentrations as permafrost thaw exposes immense stores of frozen carbon (C) to microbial decomposition. Permafrost thaw also stimulates plant growth, which could offset C loss. Using data from 7 years of experimental Air and Soil warming in moist acidic tundra, we show that Soil warming had a much stronger effect on CO₂ flux than Air warming. Soil warming caused rapid permafrost thaw and increased ecosystem respiration (R_eco), gross primary productivity (GPP), and net summer CO₂ storage (NEE). Over 7 years R_eco, GPP, and NEE also increased in Control (i.e., ambient plots), but this change could be explained by slow thaw in Control areas. In the initial stages of thaw, R_eco, GPP, and NEE increased linearly with thaw across all treatments, despite different rates of thaw. As thaw in Soil warming continued to increase linearly, ground surface subsidence created saturated microsites and suppressed R_eco, GPP, and NEE. However R_eco and GPP remained high in areas with large Eriophorum vaginatum biomass. In general NEE increased with thaw, but was more strongly correlated with plant biomass than thaw, indicating that higher R_eco in deeply thawed areas during summer months was balanced by GPP. Summer CO₂ flux across treatments fit a single quadratic relationship that captured the functional response of CO₂ flux to thaw, water table depth, and plant biomass. These results demonstrate the importance of indirect thaw effects on CO₂ flux: plant growth and water table dynamics. Nonsummer R_eco models estimated that the area was an annual CO₂ source during all years of observation. Nonsummer CO₂ loss in warmer, more deeply thawed soils exceeded the increases in summer GPP, and thawed tundra was a net annual CO₂ source.     

Environmental and Vegetation Drivers of Seasonal CO2 Fluxes in a Sub-arctic Forest–Mire Ecotone

Unravelling the role of structural and environmental drivers of gross primary productivity (GPP) and ecosystem respiration (R _eco) in highly heterogeneous tundra is a major challenge for the upscaling of chamber-based CO₂ fluxes in Arctic landscapes. In a mountain birch woodland-mire ecotone, we investigated the role of LAI (and NDVI), environmental factors (microclimate, soil moisture), and microsite type across tundra shrub plots (wet hummocks, dry hummocks, dry hollows) and lichen hummocks, in controlling net ecosystem CO₂ exchange (NEE). During a growing season, we measured NEE fluxes continuously, with closed dynamic chambers, and performed multiple fits (one for each 3-day period) of a simple light and temperature response model to hourly NEE data. Tundra shrub plots were largely CO₂ sinks, as opposed to lichen plots, although fluxes were highly variable within microsite type. For tundra shrub plots, microsite type did not influence photosynthetic parameters but it affected basal (that is, temperature-normalized) ecosystem respiration (R ₀). PAR-normalized photosynthesis (P ₆₀₀) increased with air temperature and declined with increasing vapor pressure deficit. R ₀ declined with soil moisture and showed an apparent increase with temperature, which may underlie a tight link between GPP and R _eco. NDVI was a good proxy for LAI, maximum P ₆₀₀ and maximum R ₀ of shrub plots. Cumulative CO₂ fluxes were strongly correlated with LAI (NDVI) but we observed a comparatively low GPP/LAI in dry hummocks. Our results broadly agree with the reported functional convergence across tundra vegetation, but here we show that the role of decreased productivity in transition zones and the influence of temperature and water balance on seasonal CO₂ fluxes in sub-Arctic forest–mire ecotones cannot be overlooked.     

Forest carbon balance under elevated CO2

Free-air CO₂ enrichment (FACE) technology was used to expose a loblolly pine (Pinus taeda L.) forest to elevated atmospheric CO₂ (ambient + 200 µl l^-1). After 4 years, basal area of pine trees was 9.2% larger in elevated than in ambient CO₂ plots. During the first 3 years the growth rate of pine was stimulated by ~26%. In the fourth year this stimulation declined to 23%. The average net ecosystem production (NEP) in the ambient plots was 428 gC m^-2 year^-1, indicating that the forest was a net sink for atmospheric CO₂. Elevated atmospheric CO₂ stimulated NEP by 41%. This increase was primarily an increase in plant biomass increment (57%), and secondarily increased accumulation of carbon in the forest floor (35%) and fine root increment (8%). Net primary production (NPP) was stimulated by 27%, driven primarily by increases in the growth rate of the pines. Total heterotrophic respiration (R_h) increased by 165%, but total autotrophic respiration (R_a) was unaffected. Gross primary production was increased by 18%. The largest uncertainties in the carbon budget remain in separating belowground heterotrophic (soil microbes) and autotrophic (root) respiration. If applied to temperate forests globally, the increase in NEP that we measured would fix less than 10% of the anthropogenic CO₂ projected to be released into the atmosphere in the year 2050. This may represent an upper limit because rising global temperatures, land disturbance, and heterotrophic decomposition of woody tissues will ultimately cause an increased flux of carbon back to the atmosphere.     

Effects of Warming and Clipping on Ecosystem Carbon Fluxes across Two Hydrologically Contrasting Years in an Alpine Meadow of the Qinghai-Tibet Plateau

Responses of ecosystem carbon (C) fluxes to human disturbance and climatic warming will affect terrestrial ecosystem C storage and feedback to climate change. We conducted a manipulative experiment to investigate the effects of warming and clipping on soil respiration (Rs), ecosystem respiration (ER), net ecosystem exchange (NEE) and gross ecosystem production (GEP) in an alpine meadow in a permafrost region during two hydrologically contrasting years (2012, with 29.9% higher precipitation than the long-term mean, and 2013, with 18.9% lower precipitation than the long-tem mean). Our results showed that GEP was higher than ER, leading to a net C sink (measured by NEE) over the two growing seasons. Warming significantly stimulated ecosystem C fluxes in 2012 but did not significantly affect these fluxes in 2013. On average, the warming-induced increase in GEP (1.49 µ mol m⁻²s⁻¹) was higher than in ER (0.80 µ mol m⁻²s⁻¹), resulting in an increase in NEE (0.70 µ mol m⁻²s⁻¹). Clipping and its interaction with warming had no significant effects on C fluxes, whereas clipping significantly reduced aboveground biomass (AGB) by 51.5 g m⁻² in 2013. These results suggest the response of C fluxes to warming and clipping depends on hydrological variations. In the wet year, the warming treatment caused a reduction in water, but increases in soil temperature and AGB contributed to the positive response of ecosystem C fluxes to warming. In the dry year, the reduction in soil moisture, caused by warming, and the reduction in AGB, caused by clipping, were compensated by higher soil temperatures in warmed plots. Our findings highlight the importance of changes in soil moisture in mediating the responses of ecosystem C fluxes to climate warming in an alpine meadow ecosystem.     

Contrasting above‐ and belowground organic matter decomposition and carbon and nitrogen dynamics in response to warming in High Arctic tundra

Tundra regions are projected to warm rapidly during the coming decades. The tundra biome holds the largest terrestrial carbon pool, largely contained in frozen permafrost soils. With warming, these permafrost soils may thaw and become available for microbial decomposition, potentially providing a positive feedback to global warming. Warming may directly stimulate microbial metabolism but may also indirectly stimulate organic matter turnover through increased plant productivity by soil priming from root exudates and accelerated litter turnover rates. Here, we assess the impacts of experimental warming on turnover rates of leaf litter, active layer soil and thawed permafrost sediment in two high‐arctic tundra heath sites in NE‐Greenland, either dominated by evergreen or deciduous shrubs. We incubated shrub leaf litter on the surface of control and warmed plots for 1 and 2 years. Active layer soil was collected from the plots to assess the effects of 8 years of field warming on soil carbon stocks. Finally, we incubated open cores filled with newly thawed permafrost soil for 2 years in the active layer of the same plots. After field incubation, we measured basal respiration rates of recovered thawed permafrost cores in the lab. Warming significantly reduced litter mass loss by 26% after 1 year incubation, but differences in litter mass loss among treatments disappeared after 2 years incubation. Warming also reduced litter nitrogen mineralization and decreased the litter carbon to nitrogen ratio. Active layer soil carbon stocks were reduced 15% by warming, while soil dissolved nitrogen was reduced by half in warmed plots. Warming had a positive legacy effect on carbon turnover rates in thawed permafrost cores, with 10% higher respiration rates measured in cores from warmed plots. These results demonstrate that warming may have contrasting effects on above‐ and belowground tundra carbon turnover, possibly governed by microbial resource availability.     

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.     

Unchanged carbon balance driven by equivalent responses of production and respiration to climate change in a mixed‐grass prairie

Responses of grassland carbon (C) cycling to climate change and land use remain a major uncertainty in model prediction of future climate. To explore the impacts of global change on ecosystem C fluxes and the consequent changes in C storage, we have conducted a field experiment with warming (+3 °C), altered precipitation (doubled and halved), and annual clipping at the end of growing seasons in a mixed‐grass prairie in Oklahoma, USA, from 2009 to 2013. Results showed that although ecosystem respiration (ER) and gross primary production (GPP) negatively responded to warming, net ecosystem exchange of CO₂ (NEE) did not significantly change under warming. Doubled precipitation stimulated and halved precipitation suppressed ER and GPP equivalently, with the net outcome being unchanged in NEE. These results indicate that warming and altered precipitation do not necessarily have profound impacts on ecosystem C storage. In addition, we found that clipping enhanced NEE due to a stronger positive response of GPP compared to ER, indicating that clipping could potentially be an effective land practice that could increase C storage. No significant interactions between warming, altered precipitation, and clipping were observed. Meanwhile, we found that belowground net primary production (BNPP) in general was sensitive to climate change and land use though no significant changes were found in NPP across treatments. Moreover, negative correlations of the ER/GPP ratio with soil temperature and moisture did not differ across treatments, highlighting the roles of abiotic factors in mediating ecosystem C fluxes in this grassland. Importantly, our results suggest that belowground C cycling (e.g., BNPP) could respond to climate change with no alterations in ecosystem C storage in the same period.     

CO2浓度增加和不同氮肥水平对冬小麦根系呼吸及生物量的影响

 依托FACE(Free-air CO₂ enrichment)研究平台, 利用特制分根集气生长箱, 采用静态箱-GC(Gas chromatography)法, 连续两年研究 了大气CO₂浓度升高和不同氮肥水平对冬小麦拔节期、孕穗抽穗期和灌浆末期的根系呼吸及生物量的影响。两季结果表明, CO₂浓度升高和高氮 肥量均不同程度地增加了3个阶段的地上部和地下部的生物量, 这有利于增加根茬的还田量; CO₂浓度升高对冬小麦不同生长阶段的根系呼吸影 响不同, 在拔节期影响较小;孕穗抽穗期显著增加了根系呼吸, 2004~2005季分别增加33.8%(148.1 mg N&;#8226;kg^-1 干土, HN)和43.9%(88.9 mg N&;#8226;kg^-1 干土, LN), 2005~2006季分别为23.8%(HN)和28.9%(LN); 而灌浆末期显著降低了根系呼吸, 2004~2005季分别降低31.4%(HN)和23.3% (LN), 2005~2006季分别为25.1%(HN)和18.5%(LN); 高施氮量比低施氮量促进了根系呼吸; 随着作物生长根系呼吸与地下生物量呈显著线性负相 关, 高CO₂环境中的R²变小,表明随着作物生长发育高CO₂浓度降低了作物根系呼吸与地下部生物量积累间的相关性.     

Response of tussock tundra to elevated carbon dioxide regimes: analysis of ecosystem CO2 flux through nonlinear modeling

Summary The response of tussock tundra to elevated atmospheric concentrations of CO₂ was measured at Toolik Lake, Alaska in the summer of 1983. Computer-controlled greenhouses were used to determine diurnal ecosystem flux of CO₂ under four treatments: 340 ppm, 510 ppm, and 680 ppm CO₂, as well as 680 ppm CO₂ with a four degree centrigrade increase in temperature. For the seven days of data analyzed, net daily CO₂ flux was significantly different between treatments. Net uptake was positively correlated with CO₂ concentration in the chamber and negatively correlated with temperature. A nonlinear model was used to analyze this data set and to determine some of the reasons for different net CO₂ flux. This model allowed an estimation of light utilization efficiency, total conductance of CO₂, and a comparable measure of total respiration. From this analysis we conclude that nutrient limitations in the arctic decrease the capacity of tundra plants to make use of elevated CO₂ concentrations. The plants respond by decreasing conductance in the presence of elevated CO₂, which results in approximately equal gross uptake rates for the three CO₂ treatments. Apparent changes in system respiration result in higher net uptake under elevated CO₂ but this may be due to biases in the data. The treatment with increased temperature exhibited higher conductances and, consequently, higher gross uptake of CO₂ than the other treatments. Higher temperatures, however, also increase respiration with the result being lower net uptake than would be expected in the absence of temperature inscreases.     

Belowground heathland responses after 2 years of combined warming, elevated CO2 and summer drought

Terrestrial ecosystems are exposed to atmospheric and climatic changes including increases in atmospheric CO₂ concentration, temperature and alterations of precipitation patterns, which are predicted to continue with consequences for ecosystem services and functioning in the future. In a field scale experiment on temperate heathland, manipulation of precipitation and temperature was performed with retractable curtains, and atmospheric CO₂ concentration was increased by FACE. The combination of elevated CO₂ and warming was expected to affect belowground processes additively, through increased belowground sequestration of labile carbohydrates due to elevated CO₂ in combination with temperature increased process rates. Together, these changes might increase microbial activity and availability of plant nutrients. Two years after the start of the experiment, belowground processes responded significantly to the treatments. In the combined temperature and CO₂ treatment the dissolved organic nitrogen concentration decreased and the ammonium concentration increased, but this release of nutrients was not mirrored by plant parameters. Microbial biomass carbon and microbial enrichment with ¹³C and ¹⁵N (1 year after ¹³C ₂ ¹⁵ N-glycine was injected into the soil) increased in warmed plots and in elevated CO₂ plots, but not when these treatments were combined. Furthermore, drought led to an increase in Calluna biomass and total plant nitrogen pool. The full combination of warming, elevated CO₂ and periodic drought did not unambiguously express the ecosystem responses of single factors additively, which complicates predictions of ecosystem responses to multifactor climate change.     

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.     

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.