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  1. Summary Plant phenology, the timing of recurrent biological events, shows key and complex response to climate warming, with consequences for ecosystem functions and services. A key challenge for predicting plant phenology under future climates is to determine whether the phenological changes will persist with more intensive and long‐term warming.Here, we conducted a meta‐analysis of 103 experimental warming studies around the globe to investigate the responses of four phenophases – leaf‐out, first flowering, last flowering, and leaf coloring.We showed that warming advanced leaf‐out and flowering but delayed leaf coloring across herbaceous and woody plants. As the magnitude of warming increased, the response of most plant phenophases gradually leveled off for herbaceous plants, while phenology responded in proportion to warming in woody plants. We also found that the experimental effects of warming on plant phenology diminished over time across all phenophases. Specifically, the rate of changes in first flowering for herbaceous species, as well as leaf‐out and leaf coloring for woody species, decreased as the experimental duration extended.Together, these results suggest that the real‐world impact of global warming on plant phenology will diminish over time as temperatures continue to increase. 
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  2. Abstract Tundra and boreal ecosystems encompass the northern circumpolar permafrost region and are experiencing rapid environmental change with important implications for the global carbon (C) budget. We analysed multi-decadal time series containing 302 annual estimates of carbon dioxide (CO2) flux across 70 permafrost and non-permafrost ecosystems, and 672 estimates of summer CO2flux across 181 ecosystems. We find an increase in the annual CO2sink across non-permafrost ecosystems but not permafrost ecosystems, despite similar increases in summer uptake. Thus, recent non-growing-season CO2losses have substantially impacted the CO2balance of permafrost ecosystems. Furthermore, analysis of interannual variability reveals warmer summers amplify the C cycle (increase productivity and respiration) at putatively nitrogen-limited sites and at sites less reliant on summer precipitation for water use. Our findings suggest that water and nutrient availability will be important predictors of the C-cycle response of these ecosystems to future warming. 
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  3. Abstract Accelerated warming of the Arctic can affect the global climate system by thawing permafrost and exposing organic carbon in soils to decompose and release greenhouse gases into the atmosphere. We used a process-based biosphere model (DVM-DOS-TEM) designed to simulate biophysical and biogeochemical interactions between the soil, vegetation, and atmosphere. We varied soil and environmental parameters to assess the impact on cryohydrological and biogeochemical outputs in the model. We analyzed the responses of ecosystem carbon balances to permafrost thaw by running site-level simulations at two long-term tundra ecological monitoring sites in Alaska: Eight Mile Lake (EML) and Imnavait Creek Watershed (IMN), which are characterized by similar tussock tundra vegetation but differing soil drainage conditions and climate. Model outputs showed agreement with field observations at both sites for soil physical properties and ecosystem CO2fluxes. Model simulations of Net Ecosystem Exchange (NEE) showed an overestimation during the frozen season (higher CO2emissions) at EML with a mean NEE of 26.98 ± 4.83 gC/m2/month compared to observational mean of 22.01 ± 5.67 gC/m2/month, and during the fall months at IMN, with a modeled mean of 19.21 ± 7.49 gC/m2/month compared to observation mean of 11.9 ± 4.45 gC/m2/month. Our results underscore the importance of representing the impact of soil drainage conditions on the thawing of permafrost soils, particularly poorly drained soils, which will drive the magnitude of carbon released at sites across the high-latitude tundra. These findings can help improve predictions of net carbon releases from thawing permafrost, ultimately contributing to a better understanding of the impact of Arctic warming on the global climate system. 
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  4. Abstract Large stocks of soil carbon (C) and nitrogen (N) in northern permafrost soils are vulnerable to remobilization under climate change. However, there are large uncertainties in present‐day greenhouse gas (GHG) budgets. We compare bottom‐up (data‐driven upscaling and process‐based models) and top‐down (atmospheric inversion models) budgets of carbon dioxide (CO2), methane (CH4) and nitrous oxide (N2O) as well as lateral fluxes of C and N across the region over 2000–2020. Bottom‐up approaches estimate higher land‐to‐atmosphere fluxes for all GHGs. Both bottom‐up and top‐down approaches show a sink of CO2in natural ecosystems (bottom‐up: −29 (−709, 455), top‐down: −587 (−862, −312) Tg CO2‐C yr−1) and sources of CH4(bottom‐up: 38 (22, 53), top‐down: 15 (11, 18) Tg CH4‐C yr−1) and N2O (bottom‐up: 0.7 (0.1, 1.3), top‐down: 0.09 (−0.19, 0.37) Tg N2O‐N yr−1). The combined global warming potential of all three gases (GWP‐100) cannot be distinguished from neutral. Over shorter timescales (GWP‐20), the region is a net GHG source because CH4dominates the total forcing. The net CO2sink in Boreal forests and wetlands is largely offset by fires and inland water CO2emissions as well as CH4emissions from wetlands and inland waters, with a smaller contribution from N2O emissions. Priorities for future research include the representation of inland waters in process‐based models and the compilation of process‐model ensembles for CH4and N2O. Discrepancies between bottom‐up and top‐down methods call for analyses of how prior flux ensembles impact inversion budgets, more and well‐distributed in situ GHG measurements and improved resolution in upscaling techniques. 
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  5. Abstract Arctic and alpine tundra ecosystems are large reservoirs of organic carbon1,2. Climate warming may stimulate ecosystem respiration and release carbon into the atmosphere3,4. The magnitude and persistency of this stimulation and the environmental mechanisms that drive its variation remain uncertain5–7. This hampers the accuracy of global land carbon–climate feedback projections7,8. Here we synthesize 136 datasets from 56 open-top chamber in situ warming experiments located at 28 arctic and alpine tundra sites which have been running for less than 1 year up to 25 years. We show that a mean rise of 1.4 °C [confidence interval (CI) 0.9–2.0 °C] in air and 0.4 °C [CI 0.2–0.7 °C] in soil temperature results in an increase in growing season ecosystem respiration by 30% [CI 22–38%] (n = 136). Our findings indicate that the stimulation of ecosystem respiration was due to increases in both plant-related and microbial respiration (n = 9) and continued for at least 25 years (n = 136). The magnitude of the warming effects on respiration was driven by variation in warming-induced changes in local soil conditions, that is, changes in total nitrogen concentration and pH and by context-dependent spatial variation in these conditions, in particular total nitrogen concentration and the carbon:nitrogen ratio. Tundra sites with stronger nitrogen limitations and sites in which warming had stimulated plant and microbial nutrient turnover seemed particularly sensitive in their respiration response to warming. The results highlight the importance of local soil conditions and warming-induced changes therein for future climatic impacts on respiration. 
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  6. Abstract The northern permafrost region has been projected to shift from a net sink to a net source of carbon under global warming. However, estimates of the contemporary net greenhouse gas (GHG) balance and budgets of the permafrost region remain highly uncertain. Here, we construct the first comprehensive bottom‐up budgets of CO2, CH4, and N2O across the terrestrial permafrost region using databases of more than 1000 in situ flux measurements and a land cover‐based ecosystem flux upscaling approach for the period 2000–2020. Estimates indicate that the permafrost region emitted a mean annual flux of 12 (−606, 661) Tg CO2–C yr−1, 38 (22, 53) Tg CH4–C yr−1, and 0.67 (0.07, 1.3) Tg N2O–N yr−1to the atmosphere throughout the period. Thus, the region was a net source of CH4and N2O, while the CO2balance was near neutral within its large uncertainties. Undisturbed terrestrial ecosystems had a CO2sink of −340 (−836, 156) Tg CO2–C yr−1. Vertical emissions from fire disturbances and inland waters largely offset the sink in vegetated ecosystems. When including lateral fluxes for a complete GHG budget, the permafrost region was a net source of C and N, releasing 144 (−506, 826) Tg C yr−1and 3 (2, 5) Tg N yr−1. Large uncertainty ranges in these estimates point to a need for further expansion of monitoring networks, continued data synthesis efforts, and better integration of field observations, remote sensing data, and ecosystem models to constrain the contemporary net GHG budgets of the permafrost region and track their future trajectory. 
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  7. Abstract Significant progress in permafrost carbon science made over the past decades include the identification of vast permafrost carbon stocks, the development of new pan‐Arctic permafrost maps, an increase in terrestrial measurement sites for CO2and methane fluxes, and important factors affecting carbon cycling, including vegetation changes, periods of soil freezing and thawing, wildfire, and other disturbance events. Process‐based modeling studies now include key elements of permafrost carbon cycling and advances in statistical modeling and inverse modeling enhance understanding of permafrost region C budgets. By combining existing data syntheses and model outputs, the permafrost region is likely a wetland methane source and small terrestrial ecosystem CO2sink with lower net CO2uptake toward higher latitudes, excluding wildfire emissions. For 2002–2014, the strongest CO2sink was located in western Canada (median: −52 g C m−2 y−1) and smallest sinks in Alaska, Canadian tundra, and Siberian tundra (medians: −5 to −9 g C m−2 y−1). Eurasian regions had the largest median wetland methane fluxes (16–18 g CH4m−2 y−1). Quantifying the regional scale carbon balance remains challenging because of high spatial and temporal variability and relatively low density of observations. More accurate permafrost region carbon fluxes require: (a) the development of better maps characterizing wetlands and dynamics of vegetation and disturbances, including abrupt permafrost thaw; (b) the establishment of new year‐round CO2and methane flux sites in underrepresented areas; and (c) improved models that better represent important permafrost carbon cycle dynamics, including non‐growing season emissions and disturbance effects. 
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  8. Abstract The physical and chemical changes that accompany permafrost thaw directly influence the microbial communities that mediate the decomposition of formerly frozen organic matter, leading to uncertainty in permafrost–climate feedbacks. Although changes to microbial metabolism and community structure are documented following thaw, the generality of post‐thaw assembly patterns across permafrost soils of the world remains uncertain, limiting our ability to predict biogeochemistry and microbial community responses to climate change. Based on our review of the Arctic microbiome, permafrost microbiology, and community ecology, we propose thatAssembly Theoryprovides a framework to better understand thaw‐mediated microbiome changes and the implications for community function and climate feedbacks. This framework posits that the prevalence of deterministic or stochastic processes indicates whether the community is well‐suited to thrive in changing environmental conditions. We predict that on a short timescale and following high‐disturbance thaw (e.g., thermokarst), stochasticity dominates post‐thaw microbiome assembly, suggesting that functional predictions will be aided by detailed information about the microbiome. At a longer timescale and lower‐intensity disturbance (e.g., active layer deepening), deterministic processes likely dominate, making environmental parameters sufficient for predicting function. We propose that the contribution of stochastic and deterministic processes to post‐thaw microbiome assembly depends on the characteristics of the thaw disturbance, as well as characteristics of the microbial community, such as the ecological and phylogenetic breadth of functional guilds, their functional redundancy, and biotic interactions. These propagate across space and time, potentially providing a means for predicting the microbial forcing of greenhouse gas feedbacks to global climate change. 
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  9. Abstract Earth system models (ESMs) have been rapidly developed in recent decades to advance our understanding of climate change‐carbon cycle feedback. However, those models are massive in coding, require expensive computational resources, and have difficulty in diagnosing their performance. It is highly desirable to develop ESMs with modularity and effective diagnostics. Toward these goals, we implemented a matrix approach to the Community Land Model version 5 (CLM5) to represent carbon and nitrogen cycles. Specifically, we reorganized 18 balance equations each for carbon and nitrogen cycles among the 18 vegetation pools in the original CLM5 into two matrix equations. Similarly, 140 balance equations each for carbon and nitrogen cycles among the 140 soil pools were reorganized into two additional matrix equations. The vegetation carbon and nitrogen matrix equations are connected to soil matrix equations via litterfall. The matrix equations fully reproduce simulations of carbon and nitrogen dynamics by the original model. The computational cost for forwarding simulation of the CLM5 matrix model was 26% more expensive than the original model, largely due to calculation of additional diagnostic variables, but the spin‐up computational cost was significantly saved. We showed a case study on modeled soil carbon storage under two forcing data sets to illustrate the diagnostic capability that the matrix approach uniquely offers to understand simulation results of global carbon and nitrogen dynamics. The successful implementation of the matrix approach to CLM5, one of the most complex land models, demonstrates that most, if not all, the biogeochemical models can be reorganized into the matrix form to gain high modularity, effective diagnostics, and accelerated spin‐up. 
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  10. Abstract The magnitude of future emissions of greenhouse gases from the northern permafrost region depends crucially on the mineralization of soil organic carbon (SOC) that has accumulated over millennia in these perennially frozen soils. Many recent studies have used radiocarbon (14C) to quantify the release of this “old” SOC as CO2or CH4to the atmosphere or as dissolved and particulate organic carbon (DOC and POC) to surface waters. We compiled ~1,90014C measurements from 51 sites in the northern permafrost region to assess the vulnerability of thawing SOC in tundra, forest, peatland, lake, and river ecosystems. We found that growing season soil14C‐CO2emissions generally had a modern (post‐1950s) signature, but that well‐drained, oxic soils had increased CO2emissions derived from older sources following recent thaw. The age of CO2and CH4emitted from lakes depended primarily on the age and quantity of SOC in sediments and on the mode of emission, and indicated substantial losses of previously frozen SOC from actively expanding thermokarst lakes. Increased fluvial export of aged DOC and POC occurred from sites where permafrost thaw caused soil thermal erosion. There was limited evidence supporting release of previously frozen SOC as CO2, CH4, and DOC from thawing peatlands with anoxic soils. This synthesis thus suggests widespread but not universal release of permafrost SOC following thaw. We show that different definitions of “old” sources among studies hamper the comparison of vulnerability of permafrost SOC across ecosystems and disturbances. We also highlight opportunities for future14C studies in the permafrost region. 
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