Carbon uptake in Eurasian boreal forests dominates the high-latitude net ecosystem carbon budget.

Arctic-boreal landscapes are experiencing profound warming, along with changes in ecosystem moisture status and disturbance from fire. This region is of global importance in terms of carbon feedbacks to climate, yet the sign (sink or source) and magnitude of the Arctic-boreal carbon budget within recent years remains highly uncertain. Here, we provide new estimates of recent (2003-2015) vegetation gross primary productivity (GPP), ecosystem respiration (Reco ), net ecosystem CO2 exchange (NEE; Reco - GPP), and terrestrial methane (CH4 ) emissions for the Arctic-boreal zone using a satellite data-driven process-model for northern ecosystems (TCFM-Arctic), calibrated and evaluated using measurements from >60 tower eddy covariance (EC) sites. We used TCFM-Arctic to obtain daily 1-km2 flux estimates and annual carbon budgets for the pan-Arctic-boreal region. Across the domain, the model indicated an overall average NEE sink of -850 Tg CO2 -C year-1 . Eurasian boreal zones, especially those in Siberia, contributed to a majority of the net sink. In contrast, the tundra biome was relatively carbon neutral (ranging from small sink to source). Regional CH4 emissions from tundra and boreal wetlands (not accounting for aquatic CH4 ) were estimated at 35 Tg CH4 -C year-1 . Accounting for additional emissions from open water aquatic bodies and from fire, using available estimates from the literature, reduced the total regional NEE sink by 21% and shifted many far northern tundra landscapes, and some boreal forests, to a net carbon source. This assessment, based on in situ observations and models, improves our understanding of the high-latitude carbon status and also indicates a continued need for integrated site-to-regional assessments to monitor the vulnerability of these ecosystems to climate change.

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.

Earlier snowmelt may lead to late season declines in plant productivity and carbon sequestration in Arctic tundra ecosystems.

Arctic warming is affecting snow cover and soil hydrology, with consequences for carbon sequestration in tundra ecosystems. The scarcity of observations in the Arctic has limited our understanding of the impact of covarying environmental drivers on the carbon balance of tundra ecosystems. In this study, we address some of these uncertainties through a novel record of 119 site-years of summer data from eddy covariance towers representing dominant tundra vegetation types located on continuous permafrost in the Arctic. Here we found that earlier snowmelt was associated with more tundra net CO2 sequestration and higher gross primary productivity (GPP) only in June and July, but with lower net carbon sequestration and lower GPP in August. Although higher evapotranspiration (ET) can result in soil drying with the progression of the summer, we did not find significantly lower soil moisture with earlier snowmelt, nor evidence that water stress affected GPP in the late growing season. Our results suggest that the expected increased CO2 sequestration arising from Arctic warming and the associated increase in growing season length may not materialize if tundra ecosystems are not able to continue sequestering CO2 later in the season.

Substantial hysteresis in emergent temperature sensitivity of global wetland CH4 emissions.

Wetland methane (CH4) emissions ([Formula: see text]) are important in global carbon budgets and climate change assessments. Currently, [Formula: see text] projections rely on prescribed static temperature sensitivity that varies among biogeochemical models. Meta-analyses have proposed a consistent [Formula: see text] temperature dependence across spatial scales for use in models; however, site-level studies demonstrate that [Formula: see text] are often controlled by factors beyond temperature. Here, we evaluate the relationship between [Formula: see text] and temperature using observations from the FLUXNET-CH4 database. Measurements collected across the globe show substantial seasonal hysteresis between [Formula: see text] and temperature, suggesting larger [Formula: see text] sensitivity to temperature later in the frost-free season (about 77% of site-years). Results derived from a machine-learning model and several regression models highlight the importance of representing the large spatial and temporal variability within site-years and ecosystem types. Mechanistic advancements in biogeochemical model parameterization and detailed measurements in factors modulating CH4 production are thus needed to improve global CH4 budget assessments.

Snow melt stimulates ecosystem respiration in Arctic ecosystems.

Cold seasons in Arctic ecosystems are increasingly important to the annual carbon balance of these vulnerable ecosystems. Arctic winters are largely harsh and inaccessible leading historic data gaps during that time. Until recently, cold seasons have been assumed to have negligible impacts on the annual carbon balance but as data coverage increases and the Arctic warms, the cold season has been shown to account for over half of annual methane (CH4 ) emissions and can offset summer photosynthetic carbon dioxide (CO2 ) uptake. Freeze-thaw cycle dynamics play a critical role in controlling cold season CO2 and CH4 loss, but the relationship has not been extensively studied. Here, we analyze freeze-thaw processes through in situ CO2 and CH4 fluxes in conjunction with soil cores for physical structure and porewater samples for redox biogeochemistry. We find a movement of water toward freezing fronts in soil cores, leaving air spaces in soils, which allows for rapid infiltration of oxygen-rich snow melt in spring as shown by oxidized iron in porewater. The snow melt period coincides with rising ecosystem respiration and can offset up to 41% of the summer CO2 uptake. Our study highlights this important seasonal process and shows spring greenhouse gas emissions are largely due to production from respiration instead of only bursts of stored gases. Further warming is projected to result in increases of snowpack and deeper thaws, which could increase this ecosystem respiration dominate snow melt period causing larger greenhouse gas losses during spring.

Using imaging spectroscopy to detect variation in terrestrial ecosystem productivity across a water-stressed landscape.

A central challenge to understanding how climate anomalies, such as drought and heatwaves, impact the terrestrial carbon cycle, is quantification and scaling of spatial and temporal variation in ecosystem gross primary productivity (GPP). Existing empirical and model-based satellite broadband spectra-based products have been shown to miss critical variation in GPP. Here, we evaluate the potential of high spectral resolution (10 nm) shortwave (400-2,500 nm) imagery to better detect spatial and temporal variations in GPP across a range of ecosystems, including forests, grassland-savannas, wetlands, and shrublands in a water-stressed region. Estimates of GPP from eddy covariance observations were compared against airborne hyperspectral imagery, collected across California during the 2013-2014 HyspIRI airborne preparatory campaign. Observations from 19 flux towers across 23 flight campaigns (102 total image-flux tower pairs) showed GPP to be strongly correlated to a suite of spectral wavelengths and band ratios associated with foliar physiology and chemistry. A partial least squares regression (PLSR) modeling approach was then used to predict GPP with higher validation accuracy (adjusted R2  = 0.71) and low bias (0.04) compared to existing broadband approaches (e.g., adjusted R2  = 0.68 and bias = -5.71 with the Sims et al. model). Significant wavelengths contributing to the PLSR include those previously shown to coincide with Rubisco (wavelengths 1,680, 1,740, and 2,290 nm) and Vcmax (wavelengths 1,680, 1,722, 1,732, 1,760, and 2,300 nm). These results provide strong evidence that advances in satellite spectral resolution offer significant promise for improved satellite-based monitoring of GPP variability across a diverse range of terrestrial ecosystems.

Characterizing permafrost active layer dynamics and sensitivity to landscape spatial heterogeneity in Alaska.

An important feature of the Arctic is large spatial heterogeneity in active layer conditions, which is generally poorly represented by global models and can lead to large uncertainties in predicting regional ecosystem responses and climate feedbacks. In this study, we developed a spatially integrated modelling and analysis framework combining field observations, local scale (~ 50 m resolution) active layer thickness (ALT) and soil moisture maps derived from airborne low frequency (L+P-band) radar measurements, and global satellite environmental observations to investigate the ALT sensitivity to recent climate trends and landscape heterogeneity in Alaska. Modelled ALT results show good correspondence with in situ measurements in higher permafrost probability (PP ≥ 70%) areas (n = 33, R = 0.60, mean bias = 1.58 cm, RMSE = 20.32 cm), but with larger uncertainty in sporadic and discontinuous permafrost areas. The model results also reveal widespread ALT deepening since 2001, with smaller ALT increases in northern Alaska (mean trend = 0.32 ± 1.18 cm yr-1) and much larger increases (> 3 cm yr-1) across interior and southern Alaska. The positive ALT trend coincides with regional warming and a longer snow-free season (R = 0.60 ± 0.32). A spatially integrated analysis of the radar retrievals and model sensitivity simulations demonstrated that uncertainty in the spatial and vertical distribution of soil organic carbon (SOC) was the largest factor affecting modeled ALT accuracy, while soil moisture played a secondary role. Potential improvements in characterizing SOC heterogeneity, including better spatial sampling of soil conditions and advances in remote sensing of SOC and soil moisture, will enable more accurate predictions of active layer conditions and refinement of the modelling framework across a larger domain.

Tundra water budget and implications of precipitation underestimation.

Difficulties in obtaining accurate precipitation measurements have limited meaningful hydrologic assessment for over a century due to performance challenges of conventional snowfall and rainfall gauges in windy environments. Here, we compare snowfall observations and bias adjusted snowfall to end-of-winter snow accumulation measurements on the ground for 16 years (1999-2014) and assess the implication of precipitation underestimation on the water balance for a low-gradient tundra wetland near Utqiagvik (formerly Barrow), Alaska (2007-2009). In agreement with other studies, and not accounting for sublimation, conventional snowfall gauges captured 23-56% of end-of-winter snow accumulation. Once snowfall and rainfall are bias adjusted, long-term annual precipitation estimates more than double (from 123 to 274 mm), highlighting the risk of studies using conventional or unadjusted precipitation that dramatically under-represent water balance components. Applying conventional precipitation information to the water balance analysis produced consistent storage deficits (79 to 152 mm) that were all larger than the largest actual deficit (75 mm), which was observed in the unusually low rainfall summer of 2007. Year-to-year variability in adjusted rainfall (±33 mm) was larger than evapotranspiration (±13 mm). Measured interannual variability in partitioning of snow into runoff (29% in 2008 to 68% in 2009) in years with similar end-of-winter snow accumulation (180 and 164 mm, respectively) highlights the importance of the previous summer's rainfall (25 and 60 mm, respectively) on spring runoff production. Incorrect representation of precipitation can therefore have major implications for Arctic water budget descriptions that in turn can alter estimates of carbon and energy fluxes.

CO2 exchange and evapotranspiration across dryland ecosystems of southwestern North America.

Global-scale studies suggest that dryland ecosystems dominate an increasing trend in the magnitude and interannual variability of the land CO2 sink. However, such analyses are poorly constrained by measured CO2 exchange in drylands. Here we address this observation gap with eddy covariance data from 25 sites in the water-limited Southwest region of North America with observed ranges in annual precipitation of 100-1000 mm, annual temperatures of 2-25°C, and records of 3-10 years (150 site-years in total). Annual fluxes were integrated using site-specific ecohydrologic years to group precipitation with resulting ecosystem exchanges. We found a wide range of carbon sink/source function, with mean annual net ecosystem production (NEP) varying from -350 to +330 gCm-2 across sites with diverse vegetation types, contrasting with the more constant sink typically measured in mesic ecosystems. In this region, only forest-dominated sites were consistent carbon sinks. Interannual variability of NEP, gross ecosystem production (GEP), and ecosystem respiration (Reco ) was larger than for mesic regions, and half the sites switched between functioning as C sinks/C sources in wet/dry years. The sites demonstrated coherent responses of GEP and NEP to anomalies in annual evapotranspiration (ET), used here as a proxy for annually available water after hydrologic losses. Notably, GEP and Reco were negatively related to temperature, both interannually within site and spatially across sites, in contrast to positive temperature effects commonly reported for mesic ecosystems. Models based on MODIS satellite observations matched the cross-site spatial pattern in mean annual GEP but consistently underestimated mean annual ET by ~50%. Importantly, the MODIS-based models captured only 20-30% of interannual variation magnitude. These results suggest the contribution of this dryland region to variability of regional to global CO2 exchange may be up to 3-5 times larger than current estimates.

Terrestrial carbon balance in a drier world: the effects of water availability in southwestern North America.

Global modeling efforts indicate semiarid regions dominate the increasing trend and interannual variation of net CO2 exchange with the atmosphere, mainly driven by water availability. Many semiarid regions are expected to undergo climatic drying, but the impacts on net CO2 exchange are poorly understood due to limited semiarid flux observations. Here we evaluated 121 site-years of annual eddy covariance measurements of net and gross CO2 exchange (photosynthesis and respiration), precipitation, and evapotranspiration (ET) in 21 semiarid North American ecosystems with an observed range of 100 - 1000 mm in annual precipitation and records of 4-9 years each. In addition to evaluating spatial relationships among CO2 and water fluxes across sites, we separately quantified site-level temporal relationships, representing sensitivity to interannual variation. Across the climatic and ecological gradient, photosynthesis showed a saturating spatial relationship to precipitation, whereas the photosynthesis-ET relationship was linear, suggesting ET was a better proxy for water available to drive CO2 exchanges after hydrologic losses. Both photosynthesis and respiration showed similar site-level sensitivity to interannual changes in ET among the 21 ecosystems. Furthermore, these temporal relationships were not different from the spatial relationships of long-term mean CO2 exchanges with climatic ET. Consequently, a hypothetical 100-mm change in ET, whether short term or long term, was predicted to alter net ecosystem production (NEP) by 64 gCm(-2) yr(-1). Most of the unexplained NEP variability was related to persistent, site-specific function, suggesting prioritization of research on slow-changing controls. Common temporal and spatial sensitivity to water availability increases our confidence that site-level responses to interannual weather can be extrapolated for prediction of CO2 exchanges over decadal and longer timescales relevant to societal response to climate change.

Cold season emissions dominate the Arctic tundra methane budget.

Arctic terrestrial ecosystems are major global sources of methane (CH4); hence, it is important to understand the seasonal and climatic controls on CH4 emissions from these systems. Here, we report year-round CH4 emissions from Alaskan Arctic tundra eddy flux sites and regional fluxes derived from aircraft data. We find that emissions during the cold season (September to May) account for ≥ 50% of the annual CH4 flux, with the highest emissions from noninundated upland tundra. A major fraction of cold season emissions occur during the "zero curtain" period, when subsurface soil temperatures are poised near 0 °C. The zero curtain may persist longer than the growing season, and CH4 emissions are enhanced when the duration is extended by a deep thawed layer as can occur with thick snow cover. Regional scale fluxes of CH4 derived from aircraft data demonstrate the large spatial extent of late season CH4 emissions. Scaled to the circumpolar Arctic, cold season fluxes from tundra total 12 ± 5 (95% confidence interval) Tg CH4 y(-1), ∼ 25% of global emissions from extratropical wetlands, or ∼ 6% of total global wetland methane emissions. The dominance of late-season emissions, sensitivity to soil environmental conditions, and importance of dry tundra are not currently simulated in most global climate models. Because Arctic warming disproportionally impacts the cold season, our results suggest that higher cold-season CH4 emissions will result from observed and predicted increases in snow thickness, active layer depth, and soil temperature, representing important positive feedbacks on climate warming.

Potential and limitations of inferring ecosystem photosynthetic capacity from leaf functional traits.

The aim of this study was to systematically analyze the potential and limitations of using plant functional trait observations from global databases versus in situ data to improve our understanding of vegetation impacts on ecosystem functional properties (EFPs). Using ecosystem photosynthetic capacity as an example, we first provide an objective approach to derive robust EFP estimates from gross primary productivity (GPP) obtained from eddy covariance flux measurements. Second, we investigate the impact of synchronizing EFPs and plant functional traits in time and space to evaluate their relationships, and the extent to which we can benefit from global plant trait databases to explain the variability of ecosystem photosynthetic capacity. Finally, we identify a set of plant functional traits controlling ecosystem photosynthetic capacity at selected sites. Suitable estimates of the ecosystem photosynthetic capacity can be derived from light response curve of GPP responding to radiation (photosynthetically active radiation or absorbed photosynthetically active radiation). Although the effect of climate is minimized in these calculations, the estimates indicate substantial interannual variation of the photosynthetic capacity, even after removing site-years with confounding factors like disturbance such as fire events. The relationships between foliar nitrogen concentration and ecosystem photosynthetic capacity are tighter when both of the measurements are synchronized in space and time. When using multiple plant traits simultaneously as predictors for ecosystem photosynthetic capacity variation, the combination of leaf carbon to nitrogen ratio with leaf phosphorus content explains the variance of ecosystem photosynthetic capacity best (adjusted R2 = 0.55). Overall, this study provides an objective approach to identify links between leaf level traits and canopy level processes and highlights the relevance of the dynamic nature of ecosystems. Synchronizing measurements of eddy covariance fluxes and plant traits in time and space is shown to be highly relevant to better understand the importance of intra- and interspecific trait variation on ecosystem functioning.

The uncertain climate footprint of wetlands under human pressure.

Significant climate risks are associated with a positive carbon-temperature feedback in northern latitude carbon-rich ecosystems, making an accurate analysis of human impacts on the net greenhouse gas balance of wetlands a priority. Here, we provide a coherent assessment of the climate footprint of a network of wetland sites based on simultaneous and quasi-continuous ecosystem observations of CO2 and CH4 fluxes. Experimental areas are located both in natural and in managed wetlands and cover a wide range of climatic regions, ecosystem types, and management practices. Based on direct observations we predict that sustained CH4 emissions in natural ecosystems are in the long term (i.e., several centuries) typically offset by CO2 uptake, although with large spatiotemporal variability. Using a space-for-time analogy across ecological and climatic gradients, we represent the chronosequence from natural to managed conditions to quantify the "cost" of CH4 emissions for the benefit of net carbon sequestration. With a sustained pulse-response radiative forcing model, we found a significant increase in atmospheric forcing due to land management, in particular for wetland converted to cropland. Our results quantify the role of human activities on the climate footprint of northern wetlands and call for development of active mitigation strategies for managed wetlands and new guidelines of the Intergovernmental Panel on Climate Change (IPCC) accounting for both sustained CH4 emissions and cumulative CO2 exchange.

Elevated atmospheric CO2 stimulates soil fungal diversity through increased fine root production in a semiarid shrubland ecosystem.

Soil fungal communities are likely to be central in mediating microbial feedbacks to climate change through their effects on soil carbon (C) storage, nutrient cycling, and plant health. Plants often produce increased fine root biomass in response to elevated atmospheric carbon dioxide (CO2 ), but the responses of soil microbial communities are variable and uncertain, particularly in terms of species diversity. In this study, we describe the responses of the soil fungal community to free air CO2 enrichment (FACE) in a semiarid chaparral shrubland in Southern California (dominated by Adenomstoma fasciculatum) using large subunit rRNA gene sequencing. Community composition varied greatly over the landscape and responses to FACE were subtle, involving a few specific groups. Increased frequency of Sordariomycetes and Leotiomycetes, the latter including the Helotiales, a group that includes many dark septate endophytes known to associate positively with roots, was observed in the FACE plots. Fungal diversity, both in terms of richness and evenness, increased consistently in the FACE treatment, and was relatively high compared to other studies that used similar methods. Increases in diversity were observed across multiple phylogenetic levels, from genus to class, and were distributed broadly across fungal lineages. Diversity was also higher in samples collected close to (5 cm) plants compared to samples in canopy gaps (30 cm away from plants). Fungal biomass correlated well with soil organic matter (SOM) content, but patterns of diversity were correlated with fine root production rather than SOM. We conclude that the fungal community in this ecosystem is tightly linked to plant fine root production, and that future changes in the fungal community in response to elevated CO2 and other climatic changes will be primarily driven by changes in plant belowground allocation. Potential feedbacks mediated by soil fungi, such as soil C sequestration, nutrient cycling, and pathogenesis, are discussed.

Spatial variation in landscape-level CO2 and CH4 fluxes from arctic coastal tundra: influence from vegetation, wetness, and the thaw lake cycle.

Regional quantification of arctic CO2 and CH4 fluxes remains difficult due to high landscape heterogeneity coupled with a sparse measurement network. Most of the arctic coastal tundra near Barrow, Alaska is part of the thaw lake cycle, which includes current thaw lakes and a 5500-year chronosequence of vegetated thaw lake basins. However, spatial variability in carbon fluxes from these features remains grossly understudied. Here, we present an analysis of whole-ecosystem CO2 and CH4 fluxes from 20 thaw lake cycle features during the 2011 growing season. We found that the thaw lake cycle was largely responsible for spatial variation in CO2 flux, mostly due to its control on gross primary productivity (GPP). Current lakes were significant CO2 sources that varied little. Vegetated basins showed declining GPP and CO2 sink with age (R(2) = 67% and 57%, respectively). CH4 fluxes measured from a subset of 12 vegetated basins showed no relationship with age or CO2 flux components. Instead, higher CH4 fluxes were related to greater landscape wetness (R(2) = 57%) and thaw depth (additional R(2) = 28%). Spatial variation in CO2 and CH4 fluxes had good satellite remote sensing indicators, and we estimated the region to be a small CO2 sink of -4.9 ± 2.4 (SE) g C m(-2) between 11 June and 25 August, which was countered by a CH4 source of 2.1 ± 0.2 (SE) g C m(-2) . Results from our scaling exercise showed that developing or validating regional estimates based on single tower sites can result in significant bias, on average by a factor 4 for CO2 flux and 30% for CH4 flux. Although our results are specific to the Arctic Coastal Plain of Alaska, the degree of landscape-scale variability, large-scale controls on carbon exchange, and implications for regional estimation seen here likely have wide relevance to other arctic landscapes.

Growing season and spatial variations of carbon fluxes of Arctic and boreal ecosystems in Alaska (USA).

To better understand the spatial and temporal dynamics of CO2 exchange between Arctic ecosystems and the atmosphere, we synthesized CO2 flux data, measured in eight Arctic tundra and five boreal ecosystems across Alaska (USA) and identified growing season and spatial variations of the fluxes and environmental controlling factors. For the period examined, all of the boreal and seven of the eight Arctic tundra ecosystems acted as CO2 sinks during the growing season. Seasonal patterns of the CO2 fluxes were mostly determined by air temperature, except ecosystem respiration (RE) of tundra. For the tundra ecosystems, the spatial variation of gross primary productivity (GPP) and net CO2 sink strength were explained by growing season length, whereas RE increased with growing degree days. For boreal ecosystems, the spatial variation of net CO2 sink strength was mostly determined by recovery of GPP from fire disturbance. Satellite-derived leaf area index (LAI) was a better index to explain the spatial variations of GPP and NEE of the ecosystems in Alaska than were the normalized difference vegetation index (NDVI) and enhanced vegetation index (EVI). Multiple regression models using growing degree days, growing season length, and satellite-derived LAI explained much of the spatial variation in GPP and net CO2 exchange among the tundra and boreal ecosystems. The high sensitivity of the sink strength to growing season length indicated that the tundra ecosystem could increase CO2 sink strength under expected future warming, whereas ecosystem compositions associated with fire disturbance could play a major role in carbon release from boreal ecosystems.

Light-stress avoidance mechanisms in a Sphagnum-dominated wet coastal Arctic tundra ecosystem in Alaska.

The Arctic experiences a high-radiation environment in the summer with 24-hour daylight for more than two months. Damage to plants and ecosystem metabolism can be muted by overcast conditions common in much of the Arctic. However, with climate change, extreme dry years and clearer skies could lead to the risk of increased photoxidation and photoinhibition in Arctic primary producers. Mosses, which often exceed the NPP of vascular plants in Arctic areas, are often understudied. As a result, the effect of specific environmental factors, including light, on these growth forms is poorly understood. Here, we investigated net ecosystem exchange (NEE) at the ecosystem scale, net Sphagnum CO2 exchange (NSE), and photoinhibition to better understand the impact of light on carbon exchange from a moss-dominated coastal tundra ecosystem during the summer season 2006. Sphagnum photosynthesis showed photoinhibition early in the season coupled with low ecosystem NEE. However, later in the season, Sphagnum maintained a significant CO2 uptake, probably for the development of subsurface moss layers protected from strong radiation. We suggest that the compact canopy structure of Sphagnum reduces light penetration to the subsurface layers of the moss mat and thereby protects the active photosynthetic tissues from damage. This stress avoidance mechanism allowed Sphagnum to constitute a significant percentage (up to 60%) of the ecosystem net daytime CO2 uptake at the end of the growing season despite the high levels of radiation experienced.

Top-down control of microbial activity and biomass in an Arctic soil ecosystem.

Globally, soil microbes preside over vast carbon stores, and both microbial biomass and activity are known to be regulated by bottom-up controls, that is, limitation by nutrients and energy. However, there is evidence that grazing by protozoans exerts top-down controls on biomass. Here, we investigate top-down control by phage on soil microbes using an experimental site near Barrow, Alaska (71 degrees N, 157 degrees W) during the 2007 growing season. Soil measurements were taken from sites that covered a range of microtopographical features within a drained and thawed lake basin including high- and low-centred ice-wedge polygons to estimate the availability of carbon and nitrogen for microbes. Using both field and laboratory experiments, we successfully increased both microbial biomass and respiration by decreasing phage populations. The addition of carbon and nutrients to soils had no significant effects on biomass or respiration, indicating a lack of bottom-up controls. Additionally, we present the first use of tea extracts as a potent anti-phage agent in soils. Our results suggest that top-down controls, such as phage predation, are critical to regulation of microbial activities in Arctic soils.

Effects of elevated atmospheric CO2 on soil microbial biomass, activity, and diversity in a chaparral ecosystem.

This study reports the effects of long-term elevated atmospheric CO2 on root production and microbial activity, biomass, and diversity in a chaparral ecosystem in southern California. The free air CO2 enrichment (FACE) ring was located in a stand dominated by the woody shrub Adenostoma fasciculatum. Between 1995 and 2003, the FACE ring maintained an average daytime atmospheric CO2 concentration of 550 ppm. During the last two years of operation, observations were made on soil cores collected from the FACE ring and adjacent areas of chaparral with ambient CO2 levels. Root biomass roughly doubled in the FACE plot. Microbial biomass and activity were related to soil organic matter (OM) content, and so analysis of covariance was used to detect CO2 effects while controlling for variation across the landscape. Extracellular enzymatic activity (cellulase and amylase) and microbial biomass C (chloroform fumigation-extraction) increased more rapidly with OM in the FACE plot than in controls, but glucose substrate-induced respiration (SIR) rates did not. The metabolic quotient (field respiration over potential respiration) was significantly higher in FACE samples, possibly indicating that microbial respiration was less C limited under high CO2. The treatments also differed in the ratio of SIR to microbial biomass C, indicating a metabolic difference between the microbial communities. Bacterial diversity, described by 16S rRNA clone libraries, was unaffected by the CO2 treatment, but fungal biomass was stimulated. Furthermore, fungal biomass was correlated with cellulase and amylase activities, indicating that fungi were responsible for the stimulation of enzymatic activity in the FACE treatment.

A scaling approach for quantifying the net CO2 flux of the Kuparuk River Basin, Alaska.

Net CO2 flux measurements conducted during the summer and winter of 1994-96 were scaled in space and time to provide estimates of net CO2 exchange during the 1995-96 (9 May 1995-8 May 1996) annual cycle for the Kuparuk River Basin, a 9200 km2 watershed located in NE Alaska. Net CO2 flux was measured using dynamic chambers and eddy covariance in moist-acidic, nonacidic, wet-sedge, and shrub tundra, which comprise 95% of the terrestrial landscape of the Kuparuk Basin. CO2 flux data were used as input to multivariate models that calculated instantaneous and daily rates of gross primary production (GPP) and whole-ecosystem respiration (R) as a function of meteorology and ecosystem development. Net CO2 flux was scaled up to the Kuparuk Basin using a geographical information system (GIS) consisting of a vegetation map, digital terrain map, dynamic temperature and radiation fields, and the models of GPP and R. Basin-wide estimates of net CO2 exchange for the summer growing season (9 May-5 September 1995) indicate that nonacidic tundra was a net sink of -31.7 ± 21.3 GgC (1 Gg = 109 g), while shrub tundra lost 32.5 ± 6.3 GgC to the atmosphere (negative values denote net ecosystem CO2 uptake). Acidic and wet sedge tundra were in balance, and when integrated for the entire Kuparuk River Basin (including aquatic surfaces), whole basin summer net CO2 exchange was estimated to be in balance (-0.9 ± 50.3 GgC). Autumn to winter (6 September 1995-8 May 1996) estimates of net CO2 flux indicate that acidic, nonacidic, and shrub tundra landforms were all large sources of CO2 to the atmosphere (75.5 ± 8.3, 96.4 ± 11.4, and 43.3 ± 4.7 GgC for acidic, nonacidic, and shrub tundra, respectively). CO2 loss from wet sedge surfaces was not substantially different from zero, but the large losses from the other terrestrial landforms resulted in a whole basin net CO2 loss of 217.2 ± 24.1 GgC during the 1995-96 cold season. When integrated for the 1995-96 annual cycle, acidic (66.4 + 25.25 GgC), nonacidic (64.7 ± 29.2 GgC), and shrub tundra (75.8 ± 8.4 GgC) were substantial net sources of CO2 to the atmosphere, while wet sedge tundra was in balance (0.4 + 0.8 GgC). The Kuparuk River Basin as a whole was estimated to be a net CO2 source of 218.1 ± 60.6 GgC over the 1995-96 annual cycle. Compared to direct measurements of regional net CO2 flux obtained from aircraft-based eddy covariance, the scaling procedure provided realistic estimates of CO2 exchange during the summer growing season. Although winter estimates could not be assessed directly using aircraft measurements of net CO2 exchange, the estimates reported here are comparable to measured values reported in the literature. Thus, we have high confidence in the summer estimates of net CO2 exchange and reasonable confidence in the winter net CO2 flux estimates for terrestrial landforms of the Kuparuk river basin. Although there is larger uncertainty in the aquatic estimates, the small surface area of aquatic surfaces in the Kuparuk river basin (≈ 5%) presumably reduces the potential for this uncertainty to result in large errors in basin-wide CO2 flux estimates.