Kling George W.

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George W.

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  • Preprint
    Benthic community metabolism in deep and shallow Arctic lakes during 13 years of whole–lake fertilization
    ( 2015-05) Daniels, William C. ; Kling, George W. ; Giblin, Anne E.
    Benthic primary production and oxygen consumption are important components of lake biogeochemical cycling. We performed whole-lake nutrient manipulations in arctic Alaska to assess the controls of lake morphometry, nutrients, and light on benthic community metabolism. One deep, stratified lake (Lake E5) and one shallow, well-mixed lake (Lake E6) in the Alaskan Arctic were fertilized with low levels of nitrogen (56 mg N m-3 y-1) and phosphorus (8 mg P m-3 y-1) from 2001-2013. Benthic primary production was not stimulated by fertilization in either lake. In the deep lake, decreased water clarity is consistent with an increase in phytoplankton biomass during fertilization. Benthic GPP decreased by 7 - 47 mg C m-2 d-1 (not statistically significant) and benthic respiration increased from 87 ± 20 to 167 ± 9 (SE) mg C m-2 d-1. The areal hypolimnetic oxygen deficit increased by 15 mg O2 m-2 d-1 each year during the 13 years of monitoring, apparently driven by lower (more negative) benthic NEP. In the shallow lake, phytoplankton concentration did not change with fertilization. As a result, the light environment did not change and benthic GPP did not decrease. Overall the data suggest that (1) benthic algae are not nutrient limited in either the deep or shallow lake, (2) lake morphometry modulated the overall nutrient impact on benthic metabolism by controlling the response of phytoplankton, and by extension, light and organic carbon supply to the benthos, (3) year-to-year variability in light attenuation explains considerable variability in benthic GPP between lakes and years, (4) correlations between both dissolved organic carbon concentrations and light attenuation coefficients (kd) between lakes suggests a regional control on light attenuation, and (5) the dissolved oxygen concentrations in the deep experimental lake are highly sensitive to nutrient enrichment.
  • Article
    Understanding the effects of climate change via disturbance on pristine arctic lakes-multitrophic level response and recovery to a 12-yr, low-level fertilization experiment
    (Association for the Sciences of Limnology and Oceanography, 2021-08-02) Budy, Phaedra ; Pennock, Casey A. ; Giblin, Anne E. ; Luecke, Chris ; White, Daniel L. ; Kling, George W.
    Effects of climate change-driven disturbance on lake ecosystems can be subtle; indirect effects include increased nutrient loading that could impact ecosystem function. We designed a low-level fertilization experiment to mimic persistent, climate change-driven disturbances (deeper thaw, greater weathering, or thermokarst failure) delivering nutrients to arctic lakes. We measured responses of pelagic trophic levels over 12 yr in a fertilized deep lake with fish and a shallow fishless lake, compared to paired reference lakes, and monitored recovery for 6 yr. Relative to prefertilization in the deep lake, we observed a maximum pelagic response in chl a (+201%), dissolved oxygen (DO, −43%), and zooplankton biomass (+88%) during the fertilization period (2001–2012). Other responses to fertilization, such as water transparency and fish relative abundance, were delayed, but both ultimately declined. Phyto- and zooplankton biomass and community composition shifted with fertilization. The effects of fertilization were less pronounced in the paired shallow lakes, because of a natural thermokarst failure likely impacting the reference lake. In the deep lake there was (a) moderate resistance to change in ecosystem functions at all trophic levels, (b) eventual responses were often nonlinear, and (c) postfertilization recovery (return) times were most rapid at the base of the food web (2–4 yr) while higher trophic levels failed to recover after 6 yr. The timing and magnitude of responses to fertilization in these arctic lakes were similar to responses in other lakes, suggesting indirect effects of climate change that modify nutrient inputs may affect many lakes in the future.
  • Article
    Long-term reliability of the figaro TGS 2600 solid-state methane sensor under low-Arctic conditions at Toolik Lake, Alaska
    (European Geosciences Union, 2020-05-27) Eugster, Werner ; Laundre, James A. ; Eugster, Jon ; Kling, George W.
    The TGS 2600 was the first low-cost solid-state sensor that shows a response to ambient levels of CH4 (e.g., range ≈1.8–2.7 µmol mol−1). Here we present an empirical function to correct the TGS 2600 signal for temperature and (absolute) humidity effects and address the long-term reliability of two identical sensors deployed from 2012 to 2018. We assess the performance of the sensors at 30 min resolution and aggregated to weekly medians. Over the entire period the agreement between TGS-derived and reference CH4 mole fractions measured by a high-precision Los Gatos Research instrument was R2=0.42, with better results during summer (R2=0.65 in summer 2012). Using absolute instead of relative humidity for the correction of the TGS 2600 sensor signals reduced the typical deviation from the reference to less than ±0.1 µmol mol−1 over the full range of temperatures from −41 to 27 ∘C. At weekly resolution the two sensors showed a downward drift of signal voltages indicating that after 10–13 years a TGS 2600 may have reached its end of life. While the true trend in CH4 mole fractions measured by the high-quality reference instrument was 10.1 nmolmol−1yr−1 (2012–2018), part of the downward trend in sensor signal (ca. 40 %–60 %) may be due to the increase in CH4 mole fraction because the sensor voltage decreases with increasing CH4 mole fraction. Weekly median diel cycles tend to agree surprisingly well between the TGS 2600 and reference measurements during the snow-free season, but in winter the agreement is lower. We suggest developing separate functions for deducing CH4 mole fractions from TGS 2600 measurements under cold and warm conditions. We conclude that the TGS 2600 sensor can provide data of research-grade quality if it is adequately calibrated and placed in a suitable environment where cross-sensitivities to gases other than CH4 are of no concern.
  • Article
    Nitrate is an important nitrogen source for Arctic tundra plants
    (National Academy of Sciences, 2018-03-27) Liu, Xue-Yan ; Koba, Keisuke ; Koyama, Lina A. ; Hobbie, Sarah E. ; Weiss, Marissa S. ; Inagaki, Yoshiyuki ; Shaver, Gaius R. ; Giblin, Anne E. ; Hobara, Satoru ; Nadelhoffer, Knute J. ; Sommerkorn, Martin ; Rastetter, Edward B. ; Kling, George W. ; Laundre, James A. ; Yano, Yuriko ; Makabe, Akiko ; Yano, Midori ; Liu, Cong-Qiang
    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.
  • Article
    Photochemical alteration of organic carbon draining permafrost soils shifts microbial metabolic pathways and stimulates respiration
    (Nature Publishing Group, 2017-10-03) Ward, Collin P. ; Nalven, Sarah G. ; Crump, Byron C. ; Kling, George W. ; Cory, Rose M.
    In sunlit waters, photochemical alteration of dissolved organic carbon (DOC) impacts the microbial respiration of DOC to CO2. This coupled photochemical and biological degradation of DOC is especially critical for carbon budgets in the Arctic, where thawing permafrost soils increase opportunities for DOC oxidation to CO2 in surface waters, thereby reinforcing global warming. Here we show how and why sunlight exposure impacts microbial respiration of DOC draining permafrost soils. Sunlight significantly increases or decreases microbial respiration of DOC depending on whether photo-alteration produces or removes molecules that native microbial communities used prior to light exposure. Using high-resolution chemical and microbial approaches, we show that rates of DOC processing by microbes are likely governed by a combination of the abundance and lability of DOC exported from land to water and produced by photochemical processes, and the capacity and timescale that microbial communities have to adapt to metabolize photo-altered DOC.
  • Article
    Effects of long-term climate trends on the methane and CO2 exchange processes of Toolik Lake, Alaska
    (Frontiers Media, 2022-09-13) Eugster, Werner ; DelSontro, Tonya ; Laundre, James A. ; Dobkowski, Jason ; Shaver, Gaius R. ; Kling, George W.
    Methane and carbon dioxide effluxes from aquatic systems in the Arctic will affect and likely amplify global change. As permafrost thaws in a warming world, more dissolved organic carbon (DOC) and greenhouse gases are produced and move from soils to surface waters where the DOC can be oxidized to CO2 and also released to the atmosphere. Our main study objective is to measure the release of carbon to the atmosphere via effluxes of methane (CH 4) and carbon dioxide (CO2) from Toolik Lake, a deep, dimictic, low-arctic lake in northern Alaska. By combining direct eddy covariance flux measurements with continuous gas pressure measurements in the lake surface waters, we quantified the k600 piston velocity that controls gas flux across the air–water interface. Our measured k values for CH4 and CO2 were substantially above predictions from several models at low to moderate wind speeds, and only converged on model predictions at the highest wind speeds. We attribute this higher flux at low wind speeds to effects on water-side turbulence resulting from how the surrounding tundra vegetation and topography increase atmospheric turbulence considerably in this lake, above the level observed over large ocean surfaces. We combine this process-level understanding of gas exchange with the trends of a climate-relevant long-term (30 + years) meteorological data set at Toolik Lake to examine short-term variations (2015 ice-free season) and interannual variability (2010–2015 ice-free seasons) of CH4 and CO2 fluxes. We argue that the biological processing of DOC substrate that becomes available for decomposition as the tundra soil warms is important for understanding future trends in aquatic gas fluxes, whereas the variability and long-term trends of the physical and meteorological variables primarily affect the timing of when higher or lower than average fluxes are observed. We see no evidence suggesting that a tipping point will be reached soon to change the status of the aquatic system from gas source to sink. We estimate that changes in CH4 and CO2 fluxes will be constrained with a range of +30% and −10% of their current values over the next 30 years.
  • Article
    Processing arctic eddy-flux data using a simple carbon-exchange model embedded in the ensemble Kalman filter
    (Ecological Society of America, 2010-07) Rastetter, Edward B. ; Williams, Mathew ; Griffin, Kevin L. ; Kwiatkowski, Bonnie L. ; Tomasky, Gabrielle ; Potosnak, Mark J. ; Stoy, Paul C. ; Shaver, Gaius R. ; Stieglitz, Marc ; Hobbie, John E. ; Kling, George W.
    Continuous time-series estimates of net ecosystem carbon exchange (NEE) are routinely made using eddy covariance techniques. Identifying and compensating for errors in the NEE time series can be automated using a signal processing filter like the ensemble Kalman filter (EnKF). The EnKF compares each measurement in the time series to a model prediction and updates the NEE estimate by weighting the measurement and model prediction relative to a specified measurement error estimate and an estimate of the model-prediction error that is continuously updated based on model predictions of earlier measurements in the time series. Because of the covariance among model variables, the EnKF can also update estimates of variables for which there is no direct measurement. The resulting estimates evolve through time, enabling the EnKF to be used to estimate dynamic variables like changes in leaf phenology. The evolving estimates can also serve as a means to test the embedded model and reconcile persistent deviations between observations and model predictions. We embedded a simple arctic NEE model into the EnKF and filtered data from an eddy covariance tower located in tussock tundra on the northern foothills of the Brooks Range in northern Alaska, USA. The model predicts NEE based only on leaf area, irradiance, and temperature and has been well corroborated for all the major vegetation types in the Low Arctic using chamber-based data. This is the first application of the model to eddy covariance data. We modified the EnKF by adding an adaptive noise estimator that provides a feedback between persistent model data deviations and the noise added to the ensemble of Monte Carlo simulations in the EnKF. We also ran the EnKF with both a specified leaf-area trajectory and with the EnKF sequentially recalibrating leaf-area estimates to compensate for persistent model-data deviations. When used together, adaptive noise estimation and sequential recalibration substantially improved filter performance, but it did not improve performance when used individually. The EnKF estimates of leaf area followed the expected springtime canopy phenology. However, there were also diel fluctuations in the leaf-area estimates; these are a clear indication of a model deficiency possibly related to vapor pressure effects on canopy conductance.
  • Article
    Bacterioplankton community shifts in an Arctic lake correlate with seasonal changes in organic matter source
    (American Society for Microbiology, 2003-04) Crump, Byron C. ; Kling, George W. ; Bahr, Michele ; Hobbie, John E.
    Seasonal shifts in bacterioplankton community composition in Toolik Lake, a tundra lake on the North Slope of Alaska, were related to shifts in the source (terrestrial versus phytoplankton) and lability of dissolved organic matter (DOM). A shift in community composition, measured by denaturing gradient gel electrophoresis (DGGE) of 16S rRNA genes, occurred at 4°C in near-surface waters beneath seasonal ice and snow cover in spring. This shift was associated with an annual peak in bacterial productivity ([14C]leucine incorporation) driven by the large influx of labile terrestrial DOM associated with snow meltwater. A second shift occurred after the flux of terrestrial DOM had ended in early summer as ice left the lake and as the phytoplankton community developed. Bacterioplankton communities were composed of persistent populations present throughout the year and transient populations that appeared and disappeared. Most of the transient populations could be divided into those that were advected into the lake with terrestrial DOM in spring and those that grew up from low concentrations during the development of the phytoplankton community in early summer. Sequencing of DNA in DGGE bands demonstrated that most bands represented single ribotypes and that matching bands from different samples represented identical ribotypes. Bacteria were identified as members of globally distributed freshwater phylogenetic clusters within the {alpha}- and ß-Proteobacteria, the Cytophaga-Flavobacteria-Bacteroides group, and the Actinobacteria.
  • Article
    Ecosystem responses to climate change at a Low Arctic and a High Arctic long-term research site
    (Springer, 2017-01-23) Hobbie, John E. ; Shaver, Gaius R. ; Rastetter, Edward B. ; Cherry, Jessica E. ; Goetz, Scott J. ; Guay, Kevin C. ; Gould, William A. ; Kling, George W.
    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.
  • Article
    Biomass offsets little or none of permafrost carbon release from soils, streams, and wildfire : an expert assessment
    (IOPScience, 2016-03-07) Abbott, Benjamin W. ; Jones, Jeremy B. ; Schuur, Edward A. G. ; Chapin, F. Stuart ; Bowden, William B. ; Bret-Harte, M. Syndonia ; Epstein, Howard E. ; Flannigan, Michael ; Harms, Tamara K. ; Hollingsworth, Teresa N. ; Mack, Michelle C. ; McGuire, A. David ; Natali, Susan M. ; Rocha, Adrian V. ; Tank, Suzanne E. ; Turetsky, Merritt R. ; Vonk, Jorien E. ; Wickland, Kimberly ; Aiken, George R. ; Alexander, Heather D. ; Amon, Rainer M. W. ; Benscoter, Brian ; Bergeron, Yves ; Bishop, Kevin ; Blarquez, Olivier ; Bond-Lamberty, Benjamin ; Breen, Amy L. ; Buffam, Ishi ; Cai, Yihua ; Carcaillet, Christopher ; Carey, Sean K. ; Chen, Jing M. ; Chen, Han Y. H. ; Christensen, Torben R. ; Cooper, Lee W. ; Cornelissen, Johannes H. C. ; de Groot, William J. ; DeLuca, Thomas Henry ; Dorrepaal, Ellen ; Fetcher, Ned ; Finlay, Jacques C. ; Forbes, Bruce C. ; French, Nancy H. F. ; Gauthier, Sylvie ; Girardin, Martin ; Goetz, Scott J. ; Goldammer, Johann G. ; Gough, Laura ; Grogan, Paul ; Guo, Laodong ; Higuera, Philip E. ; Hinzman, Larry ; Hu, Feng Sheng ; Hugelius, Gustaf ; JAFAROV, ELCHIN ; Jandt, Randi ; Johnstone, Jill F. ; Karlsson, Jan ; Kasischke, Eric S. ; Kattner, Gerhard ; Kelly, Ryan ; Keuper, Frida ; Kling, George W. ; Kortelainen, Pirkko ; Kouki, Jari ; Kuhry, Peter ; Laudon, Hjalmar ; Laurion, Isabelle ; Macdonald, Robie W. ; Mann, Paul J. ; Martikainen, Pertti ; McClelland, James W. ; Molau, Ulf ; Oberbauer, Steven F. ; Olefeldt, David ; Paré, David ; Parisien, Marc-André ; Payette, Serge ; Peng, Changhui ; Pokrovsky, Oleg ; Rastetter, Edward B. ; Raymond, Peter A. ; Raynolds, Martha K. ; Rein, Guillermo ; Reynolds, James F. ; Robards, Martin ; Rogers, Brendan ; Schädel, Christina ; Schaefer, Kevin ; Schmidt, Inger K. ; Shvidenko, Anatoly ; Sky, Jasper ; Spencer, Robert G. M. ; Starr, Gregory ; Striegl, Robert ; Teisserenc, Roman ; Tranvik, Lars J. ; Virtanen, Tarmo ; Welker, Jeffrey M. ; Zimov, Sergey A.
    As the permafrost region warms, its large organic carbon pool will be increasingly vulnerable to decomposition, combustion, and hydrologic export. Models predict that some portion of this release will be offset by increased production of Arctic and boreal biomass; however, the lack of robust estimates of net carbon balance increases the risk of further overshooting international emissions targets. Precise empirical or model-based assessments of the critical factors driving carbon balance are unlikely in the near future, so to address this gap, we present estimates from 98 permafrost-region experts of the response of biomass, wildfire, and hydrologic carbon flux to climate change. Results suggest that contrary to model projections, total permafrost-region biomass could decrease due to water stress and disturbance, factors that are not adequately incorporated in current models. Assessments indicate that end-of-the-century organic carbon release from Arctic rivers and collapsing coastlines could increase by 75% while carbon loss via burning could increase four-fold. Experts identified water balance, shifts in vegetation community, and permafrost degradation as the key sources of uncertainty in predicting future system response. In combination with previous findings, results suggest the permafrost region will become a carbon source to the atmosphere by 2100 regardless of warming scenario but that 65%–85% of permafrost carbon release can still be avoided if human emissions are actively reduced.
  • Article
    Arctic amplification of global warming strengthened by sunlight oxidation of permafrost carbon to CO2
    (Wiley, 2020-06-09) Bowen, Jennifer C. ; Ward, Collin P. ; Kling, George W. ; Cory, Rose M.
    Once thawed, up to 15% of the ∼1,000 Pg of organic carbon (C) in arctic permafrost soils may be oxidized to carbon dioxide (CO2) by 2,100, amplifying climate change. However, predictions of this amplification strength ignore the oxidation of permafrost C to CO2 in surface waters (photomineralization). We characterized the wavelength dependence of permafrost dissolved organic carbon (DOC) photomineralization and demonstrate that iron catalyzes photomineralization of old DOC (4,000–6,300 a BP) derived from soil lignin and tannin. Rates of CO2 production from photomineralization of permafrost DOC are twofold higher than for modern DOC. Given that model predictions of future net loss of ecosystem C from thawing permafrost do not include the loss of CO2 to the atmosphere from DOC photomineralization, current predictions of an average of 208 Pg C loss by 2,299 may be too low by ~14%.
  • Article
    Interannual, summer, and diel variability of CH4 and CO2 effluxes from Toolik Lake, Alaska, during the ice-free periods 2010-2015
    (Royal Society of Chemistry, 2020-07-09) Eugster, Werner ; DelSontro, Tonya ; Shaver, Gaius R. ; Kling, George W.
    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.
  • Article
    Biogeography of bacterioplankton in lakes and streams of an arctic tundra catchment
    (Ecological Society of America, 2007-06) Crump, Byron C. ; Adams, Heather E. ; Hobbie, John E. ; Kling, George W.
    Bacterioplankton community composition was compared across 10 lakes and 14 streams within the catchment of Toolik Lake, a tundra lake in Arctic Alaska, during seven surveys conducted over three years using denaturing gradient gel electrophoresis (DGGE) of PCR-amplified rDNA. Bacterioplankton communities in streams draining tundra were very different than those in streams draining lakes. Communities in streams draining lakes were similar to communities in lakes. In a connected series of lakes and streams, the stream communities changed with distance from the upstream lake and with changes in water chemistry, suggesting inoculation and dilution with bacteria from soil waters or hyporheic zones. In the same system, lakes shared similar bacterioplankton communities (78% similar) that shifted gradually down the catchment. In contrast, unconnected lakes contained somewhat different communities (67% similar). We found evidence that dispersal influences bacterioplankton communities via advection and dilution (mass effects) in streams, and via inoculation and subsequent growth in lakes. The spatial pattern of bacterioplankton community composition was strongly influenced by interactions among soil water, stream, and lake environments. Our results reveal large differences in lake-specific and stream-specific bacterial community composition over restricted spatial scales (<10 km) and suggest that geographic distance and connectivity influence the distribution of bacterioplankton communities across a landscape.