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We present a framework for assessing biogeochemical recovery of terrestrial ecosystems from disturbance. We identify three recovery phases. In Phase 1, nitrogen is redistributed from soil organic matter to vegetation, but the ecosystem continues to lose nitrogen because the recovering vegetation cannot take up nitrogen as fast as it is released from soil. In Phase 2, the ecosystem begins re-accumulating nitrogen and converges on a quasi-steady state in which vegetation and soil-microbial processes are in balance. In Phase 3, vegetation and soil-microbial processes remain in balance and the ecosystem slowly re-accumulates the remaining nitrogen. Phase 3 follows a balanced-accumulation trajectory along a continuum of quasi-steady states that approaches the true steady state asymptotically. We examine the effects of three ecosystem properties on recovery: openness of the nitrogen cycle, nitrogen distribution in and turnover between vegetation and soils, and the proportion of nitrogen losses that are in a refractory form. Openness exacerbates Phase 1 nitrogen losses but speeds recovery in Phases 2 and 3. A high fraction of ecosystem nitrogen in vegetation, resulting from nitrogen turnover that is slow in vegetation but fast in soil, exacerbates Phase 1 nitrogen losses but speeds recovery in Phases 2 and 3. A high proportion of nitrogen loss in refractory form mitigates Phase 1 nitrogen losses and speeds recovery in Phases 2 and 3. Application of our conceptual framework requires empirical recognition of the continuum of quasi-steady states constituting the balanced-accumulation trajectory and a distinction between the balanced-accumulation trajectory and the true steady state.

Summary • Secondary metabolites make leaves unpalatable for herbivores and influence decomposition. Site-specific differences are presented in phenolics and nitrogen in Betula nana leaves from dwarf shrub tundra at Abisko, northern Sweden, and from tussock tundra at Toolik Lake, Alaska, subjected to a decade of warming, fertilization and shading. • Nitrogen and a number of phenolics, including condensed and hydrolysable tannins, flavonoids, phenolic glucosides and chlorogenic acids, were analysed in B. nana leaves. • Phenolic concentrations showed marked between-site differences (e.g. condensed tannins were 50% higher at Abisko than at Toolik); responses to the environmental manipulations were more pronounced at Toolik compared with Abisko. Warming increased condensed tannins and decreased hydrolysable tannins at Toolik, but had no effect at Abisko, whereas fertilization and shading generally decreased concentrations. • Betula invests less carbon in phenolics at Toolik than at Abisko and shows a greater response to environmental changes by investing more carbon in growth and less to phenolic production. Hence, the Toolik population has a lower herbivore-defense level, which declines further if nutrient availability increases. By contrast, under warmer conditions, the increase in bulk phenolics and decrease in leaf palatability are greater at Toolik than at Abisko.

Principal Investigators from several sites within the Long Term Ecological Research (LTER) program offer their insights about how long-term research has been effectively sustained from periods ranging from 20 to 40 years. The sites are: Hubbard Brook (New Hampshire), Konza Prairie (Kansas), Niwot Ridge (Colorado), Arctic (Alaska), Palmer Station (Antarctica), and Georgia Coastal Ecosystems (Georgia). The main themes discussed include: the importance of a strong foundation and common vision, creating a culture of collaboration and cooperation, showing the relevance of research to societal needs, managing conflict resolution, encouraging innovation, facilitating an exchange of ideas, working to build collaborations, willingness to adopt new management structures, and careful attention to transitions in leadership. The conclusion summarizes themes based on this chapter as well as other chapters in the book.

Accelerated warming in the Arctic has led to concern regarding the amount of carbon emission potential from Arctic water bodies. Yet, aquatic carbon dioxide (CO

Accelerated warming in the Arctic has led to concern regarding the amount of carbon emission potential from Arctic water bodies. Yet, aquatic carbon dioxide (CO2) and methane (CH4) flux measurements remain scarce, particularly at high resolution and over long periods of time. Effluxes of methane (CH4) and carbon dioxide (CO2) from Toolik Lake, a deep glacial lake in northern Alaska, were measured for the first time with the direct eddy covariance (EC) flux technique during six ice-free lake periods (2010–2015). CO2 flux estimates from the lake (daily average efflux of 16.7 ± 5.3 mmol m−2 d−1) were in good agreement with earlier estimates from 1975–1989 using different methods. CH4 effluxes in 2010–2015 (averaging 0.13 ± 0.06 mmol m−2 d−1) showed an interannual variation that was 4.1 times greater than median diel variations, but mean fluxes were almost one order of magnitude lower than earlier estimates obtained from single water samples in 1990 and 2011–2012. The overall global warming potential (GWP) of Toolik Lake is thus governed mostly by CO2 effluxes, contributing 86–93% of the ice-free period GWP of 26–90 g CO2,eq m−2. Diel variation in fluxes was also important, with up to a 2-fold (CH4) to 4-fold (CO2) difference between the highest nighttime and lowest daytime effluxes. Within the summer ice-free period, on average, CH4 fluxes increased 2-fold during the first half of the summer, then remained almost constant, whereas CO2 effluxes remained almost constant over the entire summer, ending with a linear increase during the last 1–2 weeks of measurements. Due to the cold bottom temperatures of this 26 m deep lake, and the absence of ebullition and episodic flux events, Toolik Lake and other deep glacial lakes are likely not hot spots for greenhouse gas emissions, but they still contribute to the overall GWP of the Arctic.

Plant nitrogen (N) use is a key component of the N cycle in terrestrial ecosystems. The supply of N to plants affects community species composition and ecosystem processes such as photosynthesis and carbon (C) accumulation. However, the availabilities and relative importance of different N forms to plants are not well understood. While nitrate (NO3−) is a major N form used by plants worldwide, it is discounted as a N source for Arctic tundra plants because of extremely low NO3− concentrations in Arctic tundra soils, undetectable soil nitrification, and plant-tissue NO3− that is typically below detection limits. Here we reexamine NO3− use by tundra plants using a sensitive denitrifier method to analyze plant-tissue NO3−. Soil-derived NO3− was detected in tundra plant tissues, and tundra plants took up soil NO3− at comparable rates to plants from relatively NO3−-rich ecosystems in other biomes. Nitrate assimilation determined by 15N enrichments of leaf NO3− relative to soil NO3− accounted for 4 to 52% (as estimated by a Bayesian isotope-mixing model) of species-specific total leaf N of Alaskan tundra plants. Our finding that in situ soil NO3− availability for tundra plants is high has important implications for Arctic ecosystems, not only in determining species compositions, but also in determining the loss of N from soils via leaching and denitrification. Plant N uptake and soil N losses can strongly influence C uptake and accumulation in tundra soils. Accordingly, this evidence of NO3− availability in tundra soils is crucial for predicting C storage in tundra., ツンドラの生態系でも硝酸イオンは大切な窒素源だった --最先端の測定技術で「見えない」硝酸イオンの重要性を検証--. 京都大学プレスリリース. 2018-03-14.

Rapid arctic vegetation change as a result of global warming includes an increase in the cover and biomass of deciduous shrubs. Increases in shrub abundance will result in a proportional increase of shrub litter in the litter community, potentially affecting carbon turnover rates in arctic ecosystems. We investigated the effects of leaf and root litter of a deciduous shrub, Betula nana, on decomposition, by examining species-specific decomposition patterns, as well as effects of Betula litter on the decomposition of other species. We conducted a 2-yr decomposition experiment in moist acidic tundra in northern Alaska, where we decomposed three tundra species (Vaccinium vitis-idaea, Rhododendron palustre, and Eriophorum vaginatum) alone and in combination with Betula litter. Decomposition patterns for leaf and root litter were determined using three different measures of decomposition (mass loss, respiration, extracellular enzyme activity). We report faster decomposition of Betula leaf litter compared to other species, with support for species differences coming from all three measures of decomposition. Mixing effects were less consistent among the measures, with negative mixing effects shown only for mass loss. In contrast, there were few species differences or mixing effects for root decomposition. Overall, we attribute longer-term litter mass loss patterns to patterns created by early decomposition processes in the first winter. We note numerous differences for species patterns between leaf and root decomposition, indicating that conclusions from leaf litter experiments should not be extrapolated to below-ground decomposition. The high decomposition rates of Betula leaf litter aboveground, and relatively similar decomposition rates of multiple species below, suggest a potential for increases in turnover in the fast-decomposing carbon pool of leaves and fine roots as the dominance of deciduous shrubs in the Arctic increases, but this outcome may be tempered by negative litter mixing effects during the early stages of encroachment.

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.

Vegetation indices derived from solar and photosynthetically active radiation (PAR) sensors (i.e. radiation derived) have been under-utilized in inferring ecosystem function, despite measurement capability at hundreds of sites. This under-utilization may be attributed to reported mismatches among the seasonality of radiation- and satellite-derived vegetation indices and canopy photosynthesis; herein referred to as measurement biases. Here biases in radiation derived reflectance and vegetation indices were assessed using a decadal record of satellite and ground based spectroradiometer data, ecosystem phenology and CO2 fluxes, and radiation derived vegetation indices (i.e. the Normalized Difference Vegetation Index [NDVI], the two band Enhanced Vegetation Index [EVI2]) from a high latitude tundra site (i.e. Imnaviat). At Imnaviat, we found poor correspondence between the three types of reflectance and vegetation indices, especially during the latter part of the growing season. Radiation derived vegetation indices resulted in incorrect estimates of phenological timing of up to a month and poor relationships with canopy photosynthesis (i.e. Gross Ecosystem Exchange (GEE)). These mismatches were attributed to solar position (i.e. solar zenith and azimuth angle) and a method, based on the diel visible and near-infrared albedo variation, was developed to improve the performance of the vegetation indices. The ability of radiation derived vegetation indices to infer GEE and phenological dates drastically improved once radiation derived vegetation indices were corrected for solar position associated biases at Imnaviat. Moreover, radiation derived vegetation indices became better aligned with MODerate resolution Imaging Spectroradiometer (MODIS) satellite estimates after solar position associated biases were corrected at Imnaviat and at 25 Fluxnet sites (~90 site years) across North America. Corrections developed here provide a way forward in understanding daily ecosystem function or filling large gaps in eddy covariance data at a significant number of Fluxnet sites.

Rapid warming in Arctic ecosystems is resulting in increased frequency of disturbances such as fires, changes in the distribution and productivity of different plant communities, increasing thaw depths in permafrost soils and greater nutrient availability, especially nitrogen. Individually and collectively, these factors have the potential to strongly affect soil C decomposition rates, with implications for the globally significant stores of carbon in this region. However, considerable uncertainty remains regarding how C decomposition rates are controlled in Arctic soils. In this study we investigated how temperature, nitrogen availability and labile C addition affected rates of CO 2 production in short (10-day for labile C) and long-term (1.5 year for temperature and N) incubations of samples collected from burned and unburned sites in the Anaktuvuk river burn on the North Slope of Alaska from different depths (organic horizon, mineral horizon and upper permafrost). The fire in this region resulted in the loss of several cms of the organic horizon and also increased active layer depth allowing the impacts of four years of thaw on deeper soil layers to be investigated. Respiration rates did not decline substantially during the long-term incubation, although decomposition rates per unit organic matter were greater in the organic horizon. In the mineral and upper permafrost soil horizons, CO 2 production was more temperature sensitive, while N addition inhibited respiration in the mineral and upper permafrost layers, especially at low temperatures. In the short-term incubations, labile C additions promoted the decomposition of soil organic matter in the mineral and upper permafrost samples, but not in the organic samples, with this effect being lost following N addition in the deeper layers. These results highlight that (i) there are substantial amounts of labile organic matter in these soils (ii), the organic matter stored in mineral and upper permafrost in the tussock tundra is less readily decomposable than in the organic horizon, but that (iii) its decomposition is more sensitive to changes in temperature and that (iv) microbial activity in deeper soil layers is limited by labile C availability rather than N. Collectively, these results indicate that in addition to the loss of C by combustion of organic matter, increasing fire frequency also has the potential to indirectly promote the release of soil C to the atmosphere in the years following the disturbance.

A meta-analysis of soil incubation studies from the permafrost zone suggests that thawing under aerobic conditions, which releases CO2, will strengthen the permafrost carbon feedback more than waterlogged systems, which releases CO2 and CH4. Increasing temperatures in northern high latitudes are causing permafrost to thaw1, making large amounts of previously frozen organic matter vulnerable to microbial decomposition2. Permafrost thaw also creates a fragmented landscape of drier and wetter soil conditions3,4 that determine the amount and form (carbon dioxide (CO2), or methane (CH4)) of carbon (C) released to the atmosphere. The rate and form of C release control the magnitude of the permafrost C feedback, so their relative contribution with a warming climate remains unclear5,6. We quantified the effect of increasing temperature and changes from aerobic to anaerobic soil conditions using 25 soil incubation studies from the permafrost zone. Here we show, using two separate meta-analyses, that a 10 °C increase in incubation temperature increased C release by a factor of 2.0 (95% confidence interval (CI), 1.8 to 2.2). Under aerobic incubation conditions, soils released 3.4 (95% CI, 2.2 to 5.2) times more C than under anaerobic conditions. Even when accounting for the higher heat trapping capacity of CH4, soils released 2.3 (95% CI, 1.5 to 3.4) times more C under aerobic conditions. These results imply that permafrost ecosystems thawing under aerobic conditions and releasing CO2 will strengthen the permafrost C feedback more than waterlogged systems releasing CO2 and CH4 for a given amount of C.

As climate warms, changes in the carbon (C) balance of arctic tundra will play an important role in the global C balance. The C balance of tundra is tightly coupled to the nitrogen (N) and phosphorus (P) cycles because soil organic matter is the principal source of plant-available nutrients and determines the spatial variation of vegetation biomass across the North Slope of Alaska. Warming will accelerate these nutrient cycles, which should stimulate plant growth. We applied the multiple element limitation model to investigate the spatial distribution of soil organic matter and vegetation on the North Slope of Alaska and examine the effects of changes in N and P cycles on tundra C budgets under climate warming. The spatial variation of vegetation biomass on the North Slope is mainly determined by nutrient mineralization, rather than air temperature. Our simulations show substantial increases in N and P mineralization with climate warming and consequent increases in nutrient availability to plants. There are distinctly different changes in N versus P cycles in response to warming. N is lost from the region because the warming-induced increase in N mineralization is in excess of plant uptake. However, P is more tightly cycled than N and the small loss of P under warming can be compensated by entrainment of recently weathered P into the ecosystem cycle. The increase in nutrient availability results in larger C gains in vegetation than C losses from soils and hence a net accumulation of C in the ecosystems. The ongoing climate warming in Arctic enhances mineralization and leads to a net transfer of nutrient from soil organic matter to vegetation, thereby stimulating tundra plant growth and increased C sequestration in the tundra ecosystems. The C balance of the region is predominantly controlled by the internal nutrient cycles, and the external nutrient supply only exerts a minor effect on C budget.

Plants are often genetically specialized as ecotypes attuned to local environmental conditions. When conditions change, the optimal environment may be physically displaced from the local population, unless dispersal or in situ evolution keep pace, resulting in a phenomenon called adaptational lag. Using a 30-year-old reciprocal transplant study across a 475 km latitudinal gradient, we tested the adaptational lag hypothesis by measuring both short-term (tiller population growth rates) and long-term (17-year survival) fitness components of Eriophorum vaginatum ecotypes in Alaska, where climate change may have already displaced the optimum. Analyzing the transplant study as a climate transfer experiment, we showed that the climate optimum for plant performance was displaced ca. 140 km north of home sites, although plants were not generally declining in size at home sites. Adaptational lag is expected to be widespread globally for long-lived, ecotypically specialized plants, with disruptive consequences for communities and ecosystems.

Recent fire activity throughout Alaska has increased the need to understand postfire impacts on soils and permafrost vulnerability. Our study utilized data and modeling from a permafrost and ecosystem gradient to develop a mechanistic understanding of the short- and long-term impacts of tundra and boreal forest fires on soil thermal dynamics. Fires influenced a variety of factors that altered the surface energy budget, soil moisture, and the organic-layer thickness with the overall effect of increasing soil temperatures and thaw depth. The postfire thickness of the soil organic layer and its impact on soil thermal conductivity was the most important factor determining postfire soil temperatures and thaw depth. Boreal and tundra ecosystems underlain by permafrost experienced smaller postfire soil temperature increases than the nonpermafrost boreal forest from the direct and indirect effects of permafrost on drainage, soil moisture, and vegetation flammability. Permafrost decreased the loss of the insulating soil organic layer, decreased soil drying, increased surface water pooling, and created a significant heat sink to buffer postfire soil temperature and thaw depth changes. Ecosystem factors also played a role in determining postfire thaw depth with boreal forests taking several decades longer to recover their soil thermal properties than tundra. These factors resulted in tundra being less sensitive to postfire soil thermal changes than the nonpermafrost boreal forest. These results suggest that permafrost and soil organic carbon will be more vulnerable to fire as climate warms.

Moist tussock tundra, dominated by the sedge Eriophorum vaginatum L., covers approximately 3.36 × 108 km2 of arctic surface area along with large amounts of subarctic land area. Eriophorum vaginatum exhibits ecotypic differentiation along latitudinal gradients in Alaska. While ecotypic differentiation may be beneficial during periods of climate stability, it may be detrimental as climate changes, causing adaptational lag. Following harvest of a 30-year reciprocal transplant experiment, age-specific demographic data on E. vaginatum tillers were collected to parameterize a Leslie matrix. Yellow Taxi analysis, based on Tukey’s Jackknife, was used to determine mean pseudovalues of tiller population growth rate ( $$\overline{{\phi_{i} }}$$ ) for four source populations of E. vaginatum tussocks that were transplanted to each of three gardens along a latitudinal gradient. Source populations responded differentially along the latitudinal gradient. Survival and daughter tiller production influenced differences seen at the mid-latitude garden, and the overall tiller population performance was generally improved by northward transplanting relative to southward transplanting. A comparison of home-source $$\overline{{\phi_{i} }}$$ and away-source $$\overline{{\phi_{i} }}$$ within the same transplant garden indicates no home-site advantage. Although populations were still growing when transplanted to home-sites ( $$\overline{{\phi_{i} }}$$ = 1.056), tiller population growth rate increased as ΔGDD became more negative relative to the home site (i.e., as tussocks were transplanted north). These results imply that populations are affected by climate gradients in a manner consistent with adaptational lag. This study documenting the response of high-latitude ecotypes to climate gradients may be an indication of the possible future effects of climate shift in more southern latitudes.

Long-term measurements of ecological effects of warming are often not statistically significant because of annual variability or signal noise. These are reduced in indicators that filter or reduce the noise around the signal and allow effects of climate warming to emerge. In this way, certain indicators act as medium pass filters integrating the signal over years-to-decades. In the Alaskan Arctic, the 25-year record of warming of air temperature revealed no significant trend, yet environmental and ecological changes prove that warming is affecting the ecosystem. The useful indicators are deep permafrost temperatures, vegetation and shrub biomass, satellite measures of canopy reflectance (NDVI), and chemical measures of soil weathering. In contrast, the 18-year record in the Greenland Arctic revealed an extremely high summer air-warming of 1.3 °C/decade; the cover of some plant species increased while the cover of others decreased. Useful indicators of change are NDVI and the active layer thickness.

Despite concern about the status of carbon (C) in the Arctic tundra, there is currently little information on how plant respiration varies in response to environmental change in this region. We quantified the impact of long-term nitrogen (N) and phosphorus (P) treatments and greenhouse warming on the short-term temperature (T) response and sensitivity of leaf respiration (R), the high-T threshold of R, and associated traits in shoots of the Arctic shrub Betula nana in experimental plots at Toolik Lake, Alaska. Respiration only acclimated to greenhouse warming in plots provided with both N and P (resulting in a ~30% reduction in carbon efflux in shoots measured at 10 and 20 °C), suggesting a nutrient dependence of metabolic adjustment. Neither greenhouse nor N+P treatments impacted on the respiratory sensitivity to T (Q10 ); overall, Q10 values decreased with increasing measuring T, from ~3.0 at 5 °C to ~1.5 at 35 °C. New high-resolution measurements of R across a range of measuring Ts (25-70 °C) yielded insights into the T at which maximal rates of R occurred (Tmax ). Although growth temperature did not affect Tmax , N+P fertilization increased Tmax values ~5 °C, from 53 to 58 °C. N+P fertilized shoots exhibited greater rates of R than nonfertilized shoots, with this effect diminishing under greenhouse warming. Collectively, our results highlight the nutrient dependence of thermal acclimation of leaf R in B. nana, suggesting that the metabolic efficiency allowed via thermal acclimation may be impaired at current levels of soil nutrient availability. This finding has important implications for predicting carbon fluxes in Arctic ecosystems, particularly if soil N and P become more abundant in the future as the tundra warms.

High latitudes contain nearly half of global soil carbon, prompting interest in understanding how the Arctic terrestrial carbon balance will respond to rising temperatures. Low temperatures suppress the activity of soil biota, retarding decomposition and nitrogen release, which limits plant and microbial growth. Warming initially accelerates decomposition, increasing nitrogen availability, productivity and woody-plant dominance. However, these responses may be transitory, because coupled abiotic-biotic feedback loops that alter soil-temperature dynamics and change the structure and activity of soil communities, can develop. Here we report the results of a two-decade summer warming experiment in an Alaskan tundra ecosystem. Warming increased plant biomass and woody dominance, indirectly increased winter soil temperature, homogenized the soil trophic structure across horizons and suppressed surface-soil-decomposer activity, but did not change total soil carbon or nitrogen stocks, thereby increasing net ecosystem carbon storage. Notably, the strongest effects were in the mineral horizon, where warming increased decomposer activity and carbon stock: a 'biotic awakening' at depth.

In northern Alaska, variations in ecosystem characteristics, including plant species composition, soil pH, depth of the organic layer, and microbial activity, appear to be closely related to substrate age and landscape position. In this study, we conducted an extraction experiment on soils from Alaskan tundra ecosystems differing in substrate age and landscape position to elucidate the geochemical controls on solubility of organic C in these soils. The extraction experiment showed higher yields of phosphate-buffer-extractable (soluble) organic C (PEOC) in organic-layer soil from older sites with low soil pH than from younger sites with high soil pH. This was due to the strong relationship between the yield of PEOC and soil pH. Similar relationships were found with mineral soils. A significant correlation was also found between PEOC and extractable Al, suggesting that soluble organic matter is strongly adsorbed to Al oxides and hydroxides, presumably through organic chelation. The importance of geochemical factors in controlling PEOC yields was tested using soils from a variety of ecosystem types that had received long-term additions of P fertilizer. Yields of PEOC from organic soils from P-fertilized plots were significantly lower than yields from control plots, regardless of whether or not P fertilization had increased biomass or changed plant species composition in these plots. This indicates that organic matter adsorbed to minerals in organic soils is strongly controlled by geochemical factors.

I was committed to long-term, site-based, research long before the Arctic (ARC) Long-Term Ecological Research (LTER) site was established in 1987. Working with the LTER program since then has allowed me to reach my goals more easily than would have been possible otherwise. Because of my deep involvement in research in the LTER program, most of the examples I use in teaching now come from LTER sites. For the same reason, most of my communications with the public are about research in the LTER program. I learned the value of collaboration as a graduate student, from my earliest mentors and collaborators. Being a part of the LTER program has helped me to develop a wide array of enjoyable, comfortable, and productive collaborations. A message to students: be generous in all aspects of your research and professional life, because there is much more to be gained from generosity than there is to be lost. I helped set up the ARC site of the LTER program in 1987 and have made it the focus of my scientific career for the past 27 years. My experience with integrated, site-based, multidisciplinary ecosystem research actually began in 1972, however, when as a graduate student I worked with the US Tundra Biome Study at Barrow, Alaska (Brown et al. 1980; Hobbie 1980). The Tundra Biome Study and its umbrella organization, the International Biological Program (IBP), ended officially in 1974, but the ideas developed and lessons learned from these programs were central to the later development of the LTER program (Coleman 2010). These lessons were central to the formation of my own professional worldview; key among them was the idea that long-term approaches, including long-term, whole-ecosystem experiments, were essential to understanding distribution, regulation, and change in populations, communities, and ecosystems everywhere. My dissertation research, on root growth at the Barrow site, benefited greatly from the interactions I had with the diverse group who worked there. I finished my PhD in 1976, during a period when the need for a federally supported program of long-term, multidisciplinary, site-based ecological research was becoming increasingly clear.

To investigate the underlying mechanisms that control long-term recovery of tundra carbon (C) and nutrients after fire, we employed the Multiple Element Limitation (MEL) model to simulate 200-yr post-fire changes in the biogeochemistry of three sites along a burn severity gradient in response to increases in air temperature, CO2 concentration, nitrogen (N) deposition, and phosphorus (P) weathering rates. The simulations were conducted for severely burned, moderately burned, and unburned arctic tundra. Our simulations indicated that recovery of C balance after fire was mainly determined by the internal redistribution of nutrients among ecosystem components (controlled by air temperature), rather than the supply of nutrients from external sources (e.g., nitrogen deposition and fixation, phosphorus weathering). Increases in air temperature and atmospheric CO2 concentration resulted in (1) a net transfer of nutrient from soil organic matter to vegetation and (2) higher C : nutrient ratios in vegetation and soil organic matter. These changes led to gains in vegetation biomass C but net losses in soil organic C stocks. Under a warming climate, nutrients lost in wildfire were difficult to recover because the warming-induced acceleration in nutrient cycles caused further net nutrient loss from the system through leaching. In both burned and unburned tundra, the warming-caused acceleration in nutrient cycles and increases in ecosystem C stocks were eventually constrained by increases in soil C : nutrient ratios, which increased microbial retention of plant-available nutrients in the soil. Accelerated nutrient turnover, loss of C, and increasing soil temperatures will likely result in vegetation changes, which further regulate the long-term biogeochemical succession. Our analysis should help in the assessment of tundra C budgets and of the recovery of biogeochemical function following fire, which is in turn necessary for the maintenance of wildlife habitat and tundra vegetation.

© 2016, National Academy of Sciences. All rights reserved. The respiratory release of carbon dioxide (CO2) from soil is a major yet poorly understood flux in the global carbon cycle. Climatic warming is hypothesized to increase rates of soil respiration, potentially fueling further increases in global temperatures. However, despite considerable scientific attention in recent decades, the overall response of soil respiration to anticipated climatic warming remains unclear. We synthesize the largest global dataset to date of soil respiration, moisture, and temperature measurements, totaling >3,800 observations representing 27 temperature manipulation studies, spanning nine biomes and over 2 decades of warming. Our analysis reveals no significant differences in the temperature sensitivity of soil respiration between control and warmed plots in all biomes, with the exception of deserts and boreal forests. Thus, our data provide limited evidence of acclimation of soil respiration to experimental warming in several major biome types, contrary to the results from multiple single-site studies. Moreover, across all nondesert biomes, respiration rates with and without experimental warming follow a Gaussian response, increasing with soil temperature up to a threshold of ∼25 °C, above which respiration rates decrease with further increases in temperature. This consistent decrease in temperature sensitivity at higher temperatures demonstrates that rising global temperatures may result in regionally variable responses in soil respiration, with colder climates being considerably more responsive to increased ambient temperatures compared with warmer regions. Our analysis adds a unique cross-biome perspective on the temperature response of soil respiration, information critical to improving our mechanistic understanding of how soil carbon dynamics change with climatic warming.

Arctic and Boreal terrestrial ecosystems are important components of the climate system because they contain vast amounts of soil carbon (C). Evidence suggests that deciduous shrubs are increasing in abundance, but the implications for ecosystem C budgets remain uncertain. Using midsummer CO(2) flux data from 21 sites spanning 16° of latitude in the Arctic and Boreal biomes, we show that air temperature explains c. one-half of the variation in ecosystem respiration (ER) and that ER drives the pattern in net ecosystem CO(2) exchange across ecosystems. Woody sites were slightly stronger C sinks compared with herbaceous communities. However, woody sites with warm soils (> 10 °C) were net sources of CO(2) , whereas woody sites with cold soils (

Summary 1. Reciprocal transplant experiments designed to quantify genetic and environmental effects on phenotype are powerful tools for the study of local adaptation. For long-lived species, especially those in habitats with short growing seasons, however, the cumulative effects of many years in novel environments may be required for fitness differences and phenotypic changes to accrue. 2. We returned to two separate reciprocal transplant experiments thirty years after their initial establishment in interior Alaska to ask whether patterns of differentiation observed in the years immediately following transplant have persisted. We also asked whether earlier hypotheses about the role of plasticity in buffering against the effects of selection on foreign genotypes were supported. We censused survival and flowering in three transplant gardens created along a snowbank gradient for a dwarf shrub (Dryas octopetala) and six gardens created along a latitudinal gradient for a tussock-forming sedge (Eriophorum vaginatum). For both species, we used an analysis of variance to detect fitness advantages for plants transplanted back into their home site relative to those transplanted into foreign sites. 3. For D. octopetala, the original patterns of local adaptation observed in the decade following transplant appeared even stronger after three decades, with the complete elimination of foreign ecotypes in both fellfield and snowbed environments. For E. vaginatum ,d ifferential survival of populations was not evident 13 years after transplant, but was clearly evident 17 years later. There was no evidence that plasticity was associated with increased survival of foreign populations in novel sites for either D. octopetala or E. vaginatum. 4. Synthesis. We conclude that local adaptation can be strong, but nevertheless remain undetected or underestimated in short-term experiments. Such genetically based population differences limit the ability of plant populations to respond to a changing climate.

Nitrogen regulates the Earth’s climate system by constraining the terrestrial sink for atmospheric CO2. Proteolytic enzymes are a principal driver of the within-system cycle of soil nitrogen, yet there is little to no understanding of their response to climate change. Here, we use a single methodology to investigate potential proteolytic enzyme activity in soils from 16 global change experiments. We show that regardless of geographical location or experimental manipulation (i.e., temperature, precipitation, or both), all sites plotted along a single line relating the response ratio of potential proteolytic activity to soil moisture deficit, the difference between precipitation and evapotranspiration. In particular, warming and reductions in precipitation stimulated potential proteolytic activity in mesic sites – temperate and boreal forests, arctic tundra – whereas these manipulations suppressed potential activity in dry grasslands. This study provides a foundation for a simple representation of the impacts of climate change on a central component of the nitrogen cycle.

Temperature is increasing at unprecedented rates across most of the tundra biome. Remote-sensing data indicate that contemporary climate warming has already resulted in increased productivity over much of the Arctic, but plot-based evidence for vegetation transformation is not widespread. We analysed change in tundra vegetation surveyed between 1980 and 2010 in 158 plant communities spread across 46 locations. We found biome-wide trends of increased height of the plant canopy and maximum observed plant height for most vascular growth forms; increased abundance of litter; increased abundance of evergreen, low-growing and tall shrubs; and decreased abundance of bare ground. Intersite comparisons indicated an association between the degree of summer warming and change in vascular plant abundance, with shrubs, forbs and rushes increasing with warming. However, the association was dependent on the climate zone, the moisture regime and the presence of permafrost. Our data provide plot-scale evidence linking changes in vascular plant abundance to local summer warming in widely dispersed tundra locations across the globe. © 2012 Macmillan Publishers Limited. All rights reserved.

The US National Science Foundation—funded Long Term Ecological Research (LTER) Network supports a large (around 240) and diverse portfolio of long-term ecological experiments. Collectively, these long-term experiments have (a) provided unique insights into ecological patterns and processes, although such insight often became apparent only after many years of study; (b) influenced management and policy decisions; and (c) evolved into research platforms supporting studies and involving investigators who were not part of the original design. Furthermore, this suite of long-term experiments addresses, at the site level, all of the US National Research Council's Grand Challenges in Environmental Sciences. Despite these contributions, we argue that the scale and scope of global environmental change requires a more-coordinated multisite approach to long-term experiments. Ideally, such an approach would include a network of spatially extensive multifactor experiments, designed in collaboration with ecological mod...

Fire frequency has dramatically increased in the tundra of northern Alaska, USA, which has major implications for the carbon budget of the region and the functioning of these ecosystems, which support important wildlife species. We investigated the postfire succession of plant and soil carbon (C), nitrogen (N), and phosphorus (P) fluxes and stocks along a burn severity gradient in the 2007 Anaktuvuk River fire scar in northern Alaska. Modeling results indicated that the early regrowth of postfire tundra vegetation was limited primarily by its canopy photosynthetic potential, rather than nutrient availability, because of the initially low leaf area and relatively high inorganic N and P concentrations in soil. Our simulations indicated that the postfire recovery of tundra vegetation was sustained predominantly by the uptake of residual inorganic N (i.e., in the remaining ash), and the redistribution of N and P from soil organic matter to vegetation. Although residual nutrients in ash were higher in the severe burn than the moderate burn, the moderate burn recovered faster because of the higher remaining biomass and consequent photosynthetic potential. Residual nutrients in ash allowed both burn sites to recover and exceed the unburned site in both aboveground biomass and production five years after the fire. The investigation of interactions among postfire C, N, and P cycles has contributed to a mechanistic understanding of the response of tundra ecosystems to fire disturbance. Our study provided insight on how the trajectory of recovery of tundra from wildfire is regulated during early succession.

Primary producers form the base of food webs but also affect other ecosystem characteristics, such as habitat structure, light availability, and microclimate. Here, we examine changes caused by 5-30+ years of nutrient addition and resulting increases in net primary productivity (NPP) in tundra, streams, and lakes in northern Alaska. The Arctic provides an important opportunity to examine how ecosystems characterized by low diversity and low productivity respond to release from nutrient limitation. We review how responses of algae and plants affect light availability, perennial biotic structures available for consumers, oxygen levels, and temperature. Sometimes, responses were similar across all three ecosystems; e.g., increased NPP significantly reduced light to the substrate following fertilization. Perennial biotic structures increased in tundra and streams but not in lakes, and provided important new habitat niches for consumers as well as other producers. Oxygen and temperature responses also differed. Life history traits (e.g., longevity) of the primary producers along with the fate of detritus drove the responses and recovery. As global change persists and nutrients become more available in the Arctic and elsewhere, incorporating these factors as response variables will enable better prediction of ecosystem changes and feedbacks in this biome and others.

35 Abstract Understanding the sensitivity of tundra vegetation to climate warming is critical to forecasting future biodiversity and vegetation feedbacks to climate. In situ warming experiments accelerate climate change on a small scale to forecast responses of local plant communities. Limitations of this approach include the apparent site-specificity of results and uncertainty about the power of short-term studies to anticipate longer term change. We address these issues with a synthesis of 61 experimental warming studies, of up to 20 years duration, in tundra sites worldwide. The response of plant groups to warming often differed with ambient summer temperature, soil moisture and experimental duration. Shrubs increased with warming only where ambient temperature was high, whereas graminoids increased primarily in the coldest study sites. Linear increases in effect size over time were frequently observed. There was little indication of saturating or accelerating effects, as would be predicted if negative or positive vegetation feedbacks were common. These results indicate that tundra vegetation exhibits strong regional variation in response to warming, and that in vulnerable regions, cumulative effects of long-term warming on tundra vegetation - and associated ecosystem consequences - have the potential to be much greater than we have observed to date.

Little is known about how satellite imagery can be used to describe burn severity in tundra landscapes. The Anaktuvuk River Fire ARF in 2007 burned over 1000 km2 of tundra on the North Slope of Alaska, creating a mosaic of small 1 m2 to large >100 m2 patches that differed in burn severity. The ARF scar provided us with an ideal landscape to determine if a single-date spectral vegetation index can be used once vegetation recovery began and to independently determine how pixel size influences burn severity assessment. We determine and explore the sensitivity of several commonly used vegetation indices to variation in burn severity across the ARF scar and the influence of pixel size on the assessment and classification of tundra burn severity. We conducted field surveys of spectral reflectance at the peak of the first growing season post-fire extended assessment period at 18 field sites that ranged from high to low burn severity. In comparing single-date indices, we found that the two-band enhanced vegetation index EVI2 was highly correlated with normalized burn ratio NBR and better distinguished among three burn severity classes than both the NBR and the normalized difference vegetation index NDVI. We also show clear evidence that shortwave infrared SWIR reflectivity does not vary as a function of burn severity. By comparing a Quickbird scene 2.4 m pixels to simulated 30 and 250 m pixel scenes, we are able to confirm that while the moderate spatial resolution of the Landsat Thematic Mapper TM sensor 30 m is sufficient for mapping tundra burn severity, the coarser resolution of the Moderate Resolution Imaging Spectroradiometer MODIS sensor 250 m is not well matched to the fine scale of spatial heterogeneity in the ARF burn scar.

Fires produce land cover changes that have consequences for surface energy balance and temperature. Three eddy covariance towers were setup along a burn severity gradient (i.e. Severely, Moderately, and Unburned tundra) to determine the effect of fire and burn severity on arctic tundra surface energy exchange and temperature for three growing seasons (2008‐2010) following the 2007 Anaktuvuk River fire. The three sites were well matched before the fire, experienced similar weather, and had similar energy budget closure, indicating that the measured energy exchange differences between sites were largely attributable to burn severity. Increased burn severity resulted in decreased vegetation and moss cover, organic layer depth, and the rate of postfire vegetation recovery. Albedo and surface greenness steadily recovered with Moderately matching Unburned tundra by the third growing season. Decreased albedo increased net radiation and partly fueled increased latent and ground heat fluxes, soil temperatures, and thaw depth. Decreases in moss cover and the organic layer also influenced the ground thermal regime and increased latent heat fluxes. These changes either offset or decreased the surface warming effect from decreased albedo, resulting in a small surface warming in Severely and a small surface cooling in Moderately relative to Unburned tundra. These results indicate that fires have a significant impact on surface energy balance and highlight the importance of moss and permafrost thaw in regulating arctic surface energy exchange and temperature.

Burned landscapes present several challenges to quantifying landscape carbon balance. Fire scars are composed of a mosaic of patches that differ in burn severity, which may influence postfire carbon budgets through damage to vegetation and carbon stocks. We deployed three eddy covariance towers along a burn severity gradient (i.e., severely burned, moderately burned, and unburned tundra) to monitor postfire net ecosystem exchange of CO2 (NEE) within the large 2007 Anaktuvuk River fire scar in Alaska, USA, during the summer of 2008. Remote sensing data from the MODerate resolution Imaging Spectroradiometer (MODIS) was used to assess the spatial representativeness of the tower sites and parameterize a NEE model that was used to scale tower measurements to the landscape. The tower sites had similar vegetation and reflectance properties prior to the Anaktuvuk River fire and represented the range of surface conditions observed within the fire scar during the 2008 summer. Burn severity influenced a variety of surface properties, including residual organic matter, plant mortality, and vegetation recovery, which in turn determined postfire NEE. Carbon sequestration decreased with increased burn severity and was largely controlled by decreases in canopy photosynthesis. The MODIS two-band enhanced vegetation index (EVI2) monitored the seasonal course of surface greenness and explained 86% of the variability in NEE across the burn severity gradient. We demonstrate that understanding the relationship between burn severity, surface reflectance, and NEE is critical for estimating the overall postfire carbon balance of the Anaktuvuk River fire scar.

We scale a model of net ecosystem CO2 exchange (NEE) for tundra ecosystems and assess model performance using eddy covariance measurements at three tundra sites. The model, initially developed using instantaneous (seconds–minutes) chamber flux (~m2) observations, independently represents ecosystem respiration (ER) and gross primary production (GPP), and requires only temperature (T), photosynthetic photon flux density (I 0), and leaf area index (L) as inputs. We used a synthetic data set to parameterize the model so that available in situ observations could be used to assess the model. The model was then scaled temporally to daily resolution and spatially to about 1 km2 resolution, and predicted values of NEE, and associated input variables, were compared to observations obtained from eddy covariance measurements at three flux tower sites over several growing seasons. We compared observations to modeled NEE calculated using T and I 0 measured at the towers, and L derived from MODIS data. Cumulative NEE estimates were within 17 and 11% of instrumentation period and growing season observations, respectively. Predictions improved when one site-year experiencing anomalously dry conditions was excluded, indicating the potential importance of stomatal control on GPP and/or soil moisture on ER. Notable differences in model performance resulted from ER model formulations and differences in how L was estimated. Additional work is needed to gain better predictive ability in terms of ER and L. However, our results demonstrate the potential of this model to permit landscape scale estimates of NEE using relatively few and simple driving variables that are easily obtained via satellite remote sensing.

We examined short- and long-term nitrogen (N) dynamics and availability along an arctic hillslope in Alaska, USA, using a stable isotope of nitrogen ( 15 N), as a tracer. Tracer levels of 15 NH4 þ were sprayed once onto the tundra at six sites in four tundra types: heath (crest), tussock with high and low water flux (mid- and footslope), and wet sedge (riparian). 15 N in vegetation and soil was monitored to estimate retention and loss over a 3- year period. Nearly all 15 NH4 þ was immediately retained in the surface moss-detritus-plant layer, and .57% of the 15 N added remained in this layer at the end of the second year. Organic soil was the second largest 15 N sink. By the end of the third growing season, the moss-detritus-plant layer and organic soil combined retained � 87% of the 15 N added except at the Midslope site with high water flux, where recovery declined to 68%. At all sites, non-extractable and non- labile-N pools were the principal sinks for added 15 N in the organic soil. Hydrology played an important role in downslope movement of dissolved 15 N. Crest and Midslope with high-water-flux sites were most susceptible to 15 N losses via leaching, perhaps because of deep permeable mineral soil (crest) and high water flow (Midslope with high water flux). Late spring melt season also resulted in downslope dissolved- 15 N losses, perhaps because of an asynchrony between N release into melt water and soil immobilization capacity. We conclude that separation of the rooting zone from the strong sink for incoming N in the moss- detritus-plant layer, rapid incorporation of new N into relatively recalcitrant-soil-N pools within the rooting zone, and leaching loss from the upper hillslope would all contribute to the strong N-limitation of this ecosystem. An extended snow-free season and deeper depth of thaw under warmer climate may significantly alter current N dynamics in this arctic ecosystem.

Uptake of nitrogen (N) via root-mycorrhizal associations accounts for a significant portion of total N supply to many vascular plants. Using stable isotope ratios (δ15N) and the mass balance among N pools of plants, fungal tissues, and soils, a number of efforts have been made in recent years to quantify the flux of N from mycorrhizal fungi to host plants. Current estimates of this flux for arctic tundra ecosystems rely on the untested assumption that the δ15N of labile organic N taken up by the fungi is approximately the same as the δ15N of bulk soil. We report here hydrolysable amino acids are more depleted in 15N relative to hydrolysable ammonium and amino sugars in arctic tundra soils near Toolik Lake, Alaska, USA. We demonstrate, using a case study, that recognizing the depletion in 15N for hydrolysable amino acids (δ15N = −5.6‰ on average) would alter recent estimates of N flux between mycorrhizal fungi and host plants in an arctic tundra ecosystem.

A two band Enhanced Vegetation Index (EVI2) without the blue band reflectance has recently been developed as a proxy for the phenology, quantity, and activity of vegetation. We compared the ability of EVI2 and the more commonly used Normalized Difference Vegetation Index (NDVI) to resolve differences in surface greenness and Leaf Area Index (LAI) among three sites located along a burn severity gradient in arctic tundra. We calculated vegetation indices from solar and photosynthetically active radiation fluxes, and validated these calculations against vegetation indices from the Terra MODerate resolution Imaging Spectroradiometer (MODIS) and ground-based spectroradiometer measurements. EVI2 performed slightly better than NDVI when comparing tower derived vegetation indices to MODIS and spectroradiometer derived vegetation indices. Burn severity decreased albedo and resulted in differences in soil background reflectance among sites. Soil darkening had no effect on EVI2, but artificially increased NDVI, resulting in separate relationships between NDVI and Leaf Area Index for burned and unburned tundra. Our results indicate that EVI2 has several advantages over NDVI including the ability to resolve LAI differences for vegetation with different background soil reflectance.

Shrub encroachment into grass-dominated biomes is occurring globally due to a variety of anthropogenic activities, but the consequences for carbon (C) inputs, storage and cycling remain unclear. We studied eight North American graminoid-dominated ecosystems invaded by shrubs, from arctic tundra to Atlantic coastal dunes, to quantify patterns and controls of C inputs via aboveground net primary production (ANPP). Across a fourfold range in mean annual precipitation (MAP), a key regulator of ecosystem C input at the continental scale, shrub invasion decreased ANPP in xeric sites, but dramatically increased ANPP (41000gm � 2 ) at high MAP, where shrub patches maintained extraordinarily high leaf area. Concurrently, the relationship between MAP and ANPP shifted from being nonlinear in grasslands to linear in shrublands. Thus, relatively abrupt (o50 years) shifts in growth form dominance, without changes in resource quantity, can fundamentally alter continental-scale pattern of C inputs and their control by MAP in ways that exceed the direct effects of climate change alone.

Nutrient availability in the arctic is expected to increase in the next century due to accelerated decomposition associated with warming and, to a lesser extent, increased nitrogen deposition. To explore how changes in nutrient availability affect ecosystem carbon (C) cycling, we used radiocarbon to quantify changes in belowground C dynamics associated with long-term fertilization of graminoid-dominated tussock tundra at Toolik Lake, Alaska. Since 1981, yearly fertilization with nitrogen (N) and phosphorus (P) has resulted in a shift to shrub-dominated vegetation. These combined changes have altered the quantity and quality of litter inputs, the vertical distribution and dynamics of fine roots, and the decomposition rate of soil organic C. The loss of C from the deep organic and mineral soil has more than offset the C accumulation in the litter and upper organic soil horizons. In the litter and upper organic horizons, radiocarbon measurements show that increased inputs resulted in overall C accumulation, despite being offset by increased decomposition in some soil pools. To reconcile radiocarbon observations in the deeper organic and mineral soil layers, where most of the ecosystem C loss occurred, both a decrease in input of new root material and a dramatic increase of decomposition rates in centuries-old soil C pools were required. Therefore, with future increases in nutrient availability, we may expect substantial losses of C which took centuries to accumulate. © 2007 Springer Science+Business Media, LLC.

Atmospheric nitrogen (N) fixation is a key N input to arctic ecosystems, but relatively few estimates of annual N-fixation rates are available. We measured N-fixation of plant-soil cores by the acetylene reduction technique at different topographic positions in an upland tundra watershed, Imnavait Creek, through two growing seasons in order to evaluate spatial and temporal variation in N-fixation. We also examined the effects of light and temperature on N-fixation to estimate annual N-fixation rates of surface soil in this watershed using field meteorological data. Surface soil at Imnavait Creek had significant acetylene reduction potential throughout the watershed (generally 6 to 10 μmol C2H4 m−2 h−1), indicating that N-fixing organisms were present everywhere. Although acetylene reduction potential was roughly constant through the growing season, moisture, temperature and light intensity strongly affected the measured acetylene reduction rates in laboratory incubations. In addition, the relativ...

Summary 1 Northern ecosystems may lose large amounts of soil C as the global climate warms over the next few decades. This study describes how soil C loss is related to temperature, moisture and chemical composition of organic matter in Alaskan tundra soils, including soils that were fertilized annually for 8 years prior to the study. 2 Fertilized and unfertilized soils from four vegetation types (tussock, intertussock, sedge and heath) were incubated at 7 or 15 °C, under saturated or well-drained conditions, through four 100-day ‘seasons’ separated by 25- to 45-day frozen periods. 3 Losses of CO2 were monitored and total C loss was determined by difference between initial and final C stocks. Initial and final organic matter composition was determined by separation into non-polar extractable (NPE, mainly fats, oils, and waxes), water-soluble (WS, mainly soluble carbohydrates and phenolics), acid-soluble (AS, mainly cellulose and related compounds), and acid-insoluble (AIS, ‘lignin’) fractions. An isotopic label (99%13C-enriched glucose) was added to track transformations among the C fractions. 4 Total C loss during the experiment was 3–32% of initial C mass depending on soil type and treatment, with most of the loss as CO2. Wet sedge tundra soils, with the largest AS and AIS fractions, lost the least CO2 and total C. The added 13C ended up in all C fractions, indicating production, as well as loss of all fractions. 5 The greatest CO2 and total C losses occurred under warm, well-drained conditions, in all soils. The effects of fertilizer treatment were occasionally significant but never large relative to the other treatments. 6 Despite the long incubation under standard conditions, there was no evidence for convergence in C chemistry among soils as indicated by changes in relative abundances of the four C fractions. 7 Large and constant rates of C loss even after 4 ‘seasons’ of incubation suggest that a large portion of the C pool is potentially mineralizable in all soil types. 8 Warming of the Arctic climate and associated thawing of permafrost and the increase in soil drainage have the potential to cause a large release of C. This C, currently stored in soil organic matter, will be released to the atmosphere, creating a positive feedback on future climate changes.

Author Posting. © The Authors, 2005. This is the author's version of the work. It is posted here by permission of Blackwell Publishing for personal use, not for redistribution. The definitive version was published in Journal of Ecology 93 (2005): 770-782, doi:10.1111/j.1365-2745.2005.01006.x.

We used a simple model of carbon-nitrogen (C-N) interactions in terrestrial ecosystems to examine the responses to elevated CO2 and to elevated CO2 plus warming in ecosystems that had the same total nitrogen loss but that differed in the ratio of dissolved organic nitrogen (DON) to dissolved inorganic nitrogen (DIN) loss. We postulate that DIN losses can be curtailed by higher N demand in response to elevated CO2, but that DON losses cannot. We also examined simulations in which DON losses were held constant, were proportional to the amount of soil organic matter, were proportional to the soil C:N ratio, or were proportional to the rate of decomposition. We found that the mode of N loss made little difference to the short-term (,60 years) rate of carbon sequestration by the ecosystem, but high DON losses resulted in much lower carbon sequestration in the long term than did low DON losses. In the short term, C sequestration was fueled by an internal redistribution of N from soils to vegetation and by increases in the C:N ratio of soils and vegetation. This sequestration was about three times larger with elevated CO 2 and warming than with elevated CO2 alone. After year 60, C sequestration was fueled by a net accu- mulation of N in the ecosystem, and the rate of sequestration was about the same with elevated CO2 and warming as with elevated CO2 alone. With high DON losses, the ecosystem either sequestered C slowly after year 60 (when DON losses were constant or proportional to soil organic matter) or lost C (when DON losses were proportional to the soil C:N ratio or to decomposition). We conclude that changes in long-term C sequestration depend not only on the magnitude of N losses, but also on the form of those losses.

In Alaska, evergreen and deciduous shrubs dominate the vegetation of moist acidic arctic tundra (soil pH 5.5). In this study we compare soil concentrations and microbial and plant uptake of amino acids, ammonium (NH4+), and nitrate (NO3−) in acidic and nonacidic tundra. The objective was to determine any differences between the tundra sites that may relate to the differences in vegetation. We sampled the water-extractable soil N pool over one growing season and found that it at all times was higher at the nonacidic than at the acidic site, while at both sites it was dominated by NH4+ followed in order by amino acid N and NO3−. In addition, we designed an experiment in which a mixture of aspartic acid, glycine, NH4+, and NO3− were injected into the soil in the middle of the growth period. In the mixture, one N form at a time was labeled with 15N and in the case of amino acids also with 13C. Soi...

Long-term ecosystem-level experiments, in which the environment is manipulated in a controlled manner, are important tools to predict the responses of ecosystem functioning and composition to future global change. We present the results of a meta-analysis performed on the results of long-term ecosystem-level experiments near Toolik Lake, Alaska, and Abisko, Sweden. We quantified aboveground biomass responses of different arctic and subarctic ecosystems to experimental fertilization, warming and shading. We not only analysed the general patterns but also the differences in responsiveness between sites and regions. Aboveground plant biomass showed a broad similarity of responses in both locations, and also showed some important differences. In both locations, aboveground plant biomass, particularly the biomass of deciduous and graminoid plants, responded most strongly to nutrient addition. The biomass of mosses and lichens decreased in both locations as the biomass of vascular plants increased. An important difference between the two regions was the smaller positive aboveground biomass response of deciduous shrubs in Abisko as compared with Toolik Lake. Whereas in Toolik Lake Betula nana increased its dominance and replaced many of the other plant types, in Abisko all vascular plant types increased in abundance without major shifts in relative abundance. The differences between the responses of the dominant vegetation types of the Toolik Lake region, i.e. tussock tundra systems, and that of the Abisko region, i.e. heath systems, may have important implications for ecosystem development under expected patterns of global change. However, there were also large site-specific differences within each region. Several potential mechanistic explanations for the differences between sites and regions are discussed. The response patterns show the need for analyses of joint data sets from many regions and sites, in order to uncover common responses to changes in climate across large arctic regions from regional or local responses.

This study explores the relationship between the normalized difference vegetation index (NDVI), aboveground plant biomass, and ecosystem C fluxes including gross ecosystem production (GEP), ecosystem respiration (ER) and net ecosystem production. We measured NDVI across long-term experimental treatments in wet sedge tundra at the Toolik Lake LTER site, in northern Alaska. Over 13 years, N and P were applied in factorial experiments (N, P and N + P), air temperature was increased using greenhouses with and without N + P fertilizer, and light intensity (photosynthetically active photon flux density) was reduced by 50% using shade cloth. Within each treatment plot, NDVI, aboveground biomass and whole-system CO(2) flux measurements were made at the same sampling points during the peak-growing season of 2001. We found that across all treatments, NDVI is correlated with aboveground biomass ( r(2)=0.84), GEP ( r(2)=0.75) and ER ( r(2)=0.71), providing a basis for linking remotely sensed NDVI to aboveground biomass and ecosystem carbon flux.

Summary 1 Shrubs are among the tundra plants most responsive to environmental change. We measured primary and secondary stem growth in a retrospective analysis of ramets of three codominant shrubs (Betula nana, Salix pulchra, and Ledum palustre ssp. decumbens) exposed to long-term field treatment with greenhouses and N + P fertilizers at Toolik Lake, Alaska. 2 Ramets of Salix had the greatest primary stem growth under control conditions, because of their relatively high branching rate. Under fertilization, however, Betula produced much more primary stem growth than the other species, because axillary buds that would have grown as short shoots in control ramets were instead stimulated to produce long shoots (structural branches). There appeared to be a trade-off between allocation to length per stem segment and number of stem segments produced in a given year, for both Betula and Ledum. 3 Although secondary growth in stems is the largest component of above-ground net primary production in forests, it is often ignored in shrub-dominated ecosystems. We derived an expression for secondary growth in shrubs based on distributions of stem mass and length with age, and allowing for experimentally induced changes in secondary growth rate. 4 There was good agreement between measured ramet stem mass and calculated values for all three species, validating our mathematical analysis of secondary growth. 5 Fertilization greatly increased the relative rate of secondary growth only in Betula, consistent with observed accumulations of its stem mass in ecosystem-level quadrat harvests. Secondary growth of Betula was a major component of ecosystem NPP in fertilized plots and probably contributes significantly to ecosystem carbon storage. 6 The increase in its secondary growth enabled Betula to become dominant under fertilization, whereas the inability of older stems of Ledum to respond in this way prevented it from growing into the canopy.