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2021
Duffy KA, Schwalm CR, Arcus VL, Koch GW, Liang LL, Schipper LA (2021) How close are we to the temperature tipping point of the terrestrial biosphere?. Science Advances 7(3):eaay1052.
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Read Publication<span>The temperature dependence of global photosynthesis and respiration determine land carbon sink strength. While the land sink currently mitigates ~30% of anthropogenic carbon emissions, it is unclear whether this ecosystem service will persist and, more specifically, what hard temperature limits, if any, regulate carbon uptake. Here, we use the largest continuous carbon flux monitoring network to construct the first observationally derived temperature response curves for global land carbon uptake. We show that the mean temperature of the warmest quarter (3-month period) passed the thermal maximum for photosynthesis during the past decade. At higher temperatures, respiration rates continue to rise in contrast to sharply declining rates of photosynthesis. Under business-as-usual emissions, this divergence elicits a near halving of the land sink strength by as early as 2040.</span>
2020
Balazs KR, Kramer AT, Munson SM, Talkington N, Still S (2020) The right trait in the right place at the right time: Matching traits to environment improves restoration outcomes. Ecological Applications 30(4): e02110.
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Read Publication<span>The challenges of restoration in dryland ecosystems are growing due to a rise in anthropogenic disturbance and increasing aridity. Plant functional traits are often used to predict plant performance and can offer a window into potential outcomes of restoration efforts across environmental gradients. We analyzed a database including 15 yr of seeding outcomes across 150 sites on the Colorado Plateau, a cold desert ecoregion in the western United States, and analyzed the independent and interactive effects of functional traits (seed mass, height, and specific leaf area) and local biologically relevant climate variables on seeding success. We predicted that the best models would include an interaction between plant traits and climate, indicating a need to match the right trait value to the right climate conditions to maximize seeding success. Indeed, we found that both plant height and seed size significantly interacted with temperature seasonality, with larger seeds and taller plants performing better in more seasonal environments. We also determined that these trait?environment patterns are not influenced by whether a species is native or nonnative. Our results inform the selection of seed mixes for restoring areas with specific climatic conditions, while also demonstrating the strong influence of temperature seasonality on seeding success in the Colorado Plateau region.</span>
Birch L, Schwalm CR, Natali S, Lombardozzi D, Keppel-Aleks G, Watts J, Lin X, Zona D, Oechel W, Sachs T, Black TA, Rogers BM (2020) Addressing Biases in Arctic-Boreal Carbon Cycling in the Community Land Model Version 5. Geosci. Model Dev. Discuss. 2020: 1-31.
Read PublicationBlazewicz SJ, Hungate BA, Koch BJ, Nuccio EE, Morrissey E, Brodie EL, Schwartz E, Pett-Ridge J, Firestone MK (2020) Taxon-specific microbial growth and mortality patterns reveal distinct temporal population responses to rewetting in a California grassland soil. The ISME Journal 14(6): 1520-1532.
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Read Publication<span>Microbial activity increases after rewetting dry soil, resulting in a pulse of carbon mineralization and nutrient availability. The biogeochemical responses to wet-up are reasonably well understood and known to be microbially mediated. Yet, the population level dynamics, and the resulting changes in microbial community patterns, are not well understood as ecological phenomena. Here, we used sequencing of 16S rRNA genes coupled with heavy water (H218O) DNA quantitative stable isotope probing to estimate population-specific rates of growth and mortality in response to a simulated wet-up event in a California annual grassland soil. Bacterial growth and mortality responded rapidly to wet-up, within 3 h, and continued throughout the 168 h incubation, with patterns of sequential growth observed at the phylum level. Of the 37 phyla detected in the prewet community, growth was found in 18 phyla while mortality was measured in 26 phyla. Rapid growth and mortality rates were measurable within 3 h of wet-up but had contrasting characteristics; growth at 3 h was dominated by select taxa within the Proteobacteria and Firmicutes, whereas mortality was taxonomically widespread. Furthermore, across the community, mortality exhibited density-independence, consistent with the indiscriminate shock resulting from dry-down and wet-up, whereas growth was density-dependent, consistent with control by competition or predation. Total aggregated growth across the community was highly correlated with total soil CO2 production. Together, these results illustrate how previously “invisible” population responses can translate quantitatively to emergent observations of ecosystem-scale biogeochemistry.</span>
Blumstein M, Richardson A, Weston D, Zhang J, Muchero W, Hopkins R (2020) A new perspective on ecological prediction reveals limits to climate adaptation in a temperate tree species. Current Biology 30(8): 1447-1453.e4.
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Read Publication<span>Forests absorb a large fraction of anthropogenic CO2 emission, but their ability to continue to act as a sink under climate change depends in part on plant species undergoing rapid adaptation. Yet models of forest response to climate change currently ignore local adaptation as a response mechanism. Thus, considering the evolution of intraspecific trait variation is necessary for reliable, long-term species and climate projections. Here, we combine ecophysiology and predictive climate modeling with analyses of genomic variation to determine whether sugar and starch storage, energy reserves for trees under extreme conditions, have the heritable variation and genetic diversity necessary to evolve in response to climate change within populations of black cottonwood (Populus trichocarpa). Despite current patterns of local adaptation and extensive range-wide heritable variation in storage, we demonstrate that adaptive evolution in response to climate change will be limited by a lack of heritable variation within northern populations and by a need for extreme genetic changes in southern populations. Our method can help design more targeted species management interventions and highlights the power of using genomic tools in ecological prediction to scale from molecular to regional processes to determine the ability of a species to respond to future climates.</span>
Bond-Lamberty B, Christianson DS, Malhotra A, Pennington S, Sihi D, AghaKouchak A, Anjileli H, Altaf AM, Armesto JJ, Ashraf S, Ataka M, Baldocchi D, Andrew Black T, Buchmann N, Carbone MS,Chang SC, Crill P, Curtis PS, Davidson EA, Desai AR, Drake JE, El-Madany TS, Gavazzi M, Görres CM, Gough CM, Goulden M, Gregg J, Gutiérrez del Arroyo O, He JS, Hirano T, Hopple A, Hughes H, Järveoja J, Jassal R, Jian J, Kan H, Kaye J, Kominami Y, Liang N, Lipson D, Macdonald CA, Maseyk K, Mathes K, Mauritz M, Mayes MA, McNulty S, Miao G, Migliavacca M, Miller S, Miniat CF, Nietz JG, Nilsson MB, Moormets A, Norouzi H, O'Connell CS, Osborne B, Oyonarte C, Pang Z, Peichl M, Pendall E, Perez-Quezada JF, Phillips CL, Phillips RP, Raich JW, Renchon AA, Ruehr NK, Sanchez-Cañete EP, Saunders M, Savage KE, Schrumpf M, Scott RL, Seibt U, Silver WL, Sun W, Szutu D, Takagi K, Takagi M, Teramoto M, Tjoelker MG, Trumbore S, Ueyama M, Vargas R, Varner RK, Verfaillie J, Vogel C, Wang J, Winston G, Wood TE, Wu J, Wutzler T, Zeng J, Zha T, Zhang Q, Zou J (2020) COSORE: A community database for continuous soil respiration and other soil-atmosphere greenhouse gas flux data. Global Change Biology 26(12): 7268-7283.
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Read Publication<span>Globally, soils store two to three times as much carbon as currently resides in the atmosphere, and it is critical to understand how soil greenhouse gas (GHG) emissions and uptake will respond to ongoing climate change. In particular, the soil-to-atmosphere CO2 flux, commonly though imprecisely termed soil respiration (RS), is one of the largest carbon fluxes in the Earth system. An increasing number of high-frequency RS measurements (typically, from an automated system with hourly sampling) have been made over the last two decades; an increasing number of methane measurements are being made with such systems as well. Such high frequency data are an invaluable resource for understanding GHG fluxes, but lack a central database or repository. Here we describe the lightweight, open-source COSORE (COntinuous SOil REspiration) database and software, that focuses on automated, continuous and long-term GHG flux datasets, and is intended to serve as a community resource for earth sciences, climate change syntheses and model evaluation. Contributed datasets are mapped to a single, consistent standard, with metadata on contributors, geographic location, measurement conditions and ancillary data. The design emphasizes the importance of reproducibility, scientific transparency and open access to data. While being oriented towards continuously measured RS, the database design accommodates other soil-atmosphere measurements (e.g. ecosystem respiration, chamber-measured net ecosystem exchange, methane fluxes) as well as experimental treatments (heterotrophic only, etc.). We give brief examples of the types of analyses possible using this new community resource and describe its accompanying R software package.</span>
Butterfield BJ, Grams PE, Durning LE, Hazel J, Palmquist EC, Ralston BE, Sankey JB (2020) Associations between riparian plant morphological guilds and fluvial sediment dynamics along the regulated Colorado River in Grand Canyon. River Research and Applications 36(3): 410-421.
Read Abstract<span>Effects of riparian vegetation on fluvial sediment dynamics depend on morphological traits of the constituent species. Determining the effects of different morphological guilds on sedimentation rates, as influenced by multiple aspects of dam operations, can help identify viable strategies for streamflow and vegetation management to achieve riparian resource goals. Plants of increasing size and branching density or complexity have been found to have greater effects on sedimentation in free-flowing systems; however, this relationship could differ in regulated rivers. We tested the hypothesis that plant guilds of increasing height and branching complexity would be positively associated with sedimentation rates on 23 sandbars deposited in zones of recirculating flow (eddies) along the Colorado River in Grand Canyon. We used an image-based vegetation classification and digital elevation models from annual topographic surveys to track associations between six plant morphological guilds and topographic change over 5?years. Vegetation had significant associations with deposition after accounting for geomorphic setting, but the ordinal guild scale was not positively correlated with deposition magnitude. Instead, low-statured rhizomatous and herbaceous guilds were particularly effective at capturing sediment in the separation zone of sandbars, whereas tall herbs and large shrubs were most effective at capturing sediment in reattachment zones. These nuanced interactions between geomorphic position and morphological guild may be a direct consequence of flow regulation through modifications to physical deposition and erosion processes. Flow regulation may also select for a narrow subset of morphological guilds, reducing the diversity of vegetation feedbacks on sedimentation and emphasizing geomorphic drivers.</span>
Chen C, van Groenigen KJ, Yang H, Hungate BA, Yang B, Tian Y, Chen J, Dong W, Huang S, Deng A, Jiang Y, Zhang W (2020) Global warming and shifts in cropping systems together reduce China’s rice production. Global Food Security 24(100359).
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Read Publication<span>Climate warming is widely expected to affect rice yields, but results are equivocal and variation in rice cropping systems and climatic conditions complicates country-scale yield assessments. Here we show, through meta–analysis of field warming experiments, that yield responses to warming differ strongly between China's rice cropping systems. Whereas warming increases yields in “single rice” systems, it decreases yields in “middle rice” systems and has contrasting effects for early and late rice in “double rice” systems. We further show that the contribution of these cropping systems to China's total rice production has shifted dramatically over recent decades. We estimate that if the present structure of rice cropping systems persists, warming will reduce China's total rice production by 5.0% in 2060. However, if the recent decline in the area of double rice systems continues, China's rice production may decrease by 13.5%. Our results underline the need for maintaining the current area of China's “double rice” cropping system and for technological innovations in multiple rice cropping systems to ensure food security in a warming climate.</span>
Chen J, Elsgaard L, van Groenigen KJ, Olesen JE, Liang Z, Jiang Y, Laerke PE, Zhang Y, Luo Y, Hungate BA, Sinsabaugh RL, Jorgensen U (2020) Soil carbon loss with warming: New evidence from carbon-degrading enzymes. Global Change Biology 26(4): 1944-1952.
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Read Publication<span>Climate warming affects soil carbon (C) dynamics, with possible serious consequences for soil C stocks and atmospheric CO2 concentrations. However, the mechanisms underlying changes in soil C storage are not well understood, hampering long-term predictions of climate C-feedbacks. The activity of the extracellular enzymes ligninase and cellulase can be used to track changes in the predominant C sources of soil microbes and can thus provide mechanistic insights into soil C loss pathways. Here we show, using meta-analysis, that reductions in soil C stocks with warming are associated with increased ratios of ligninase to cellulase activity. Furthermore, whereas long-term (≥5 years) warming reduced the soil recalcitrant C pool by 14%, short-term warming had no significant effect. Together, these results suggest that warming stimulates microbial utilization of recalcitrant C pools, possibly exacerbating long-term climate-C feedbacks.</span>
Chen J, van Groenigen KJ, Hungate BA, Terrer C, van Groenigen JW, Maestre FT, Ying SC, Luo Y, Jorgensen U, Sinsabaugh RL, Olesen JE, Elsgaard L (2020) Long-term nitrogen loading alleviates phosphorus limitation in terrestrial ecosystems. Global Change Biology.
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Read Publication<span>Increased human-derived nitrogen (N) deposition to terrestrial ecosystems has resulted in widespread phosphorus (P) limitation of net primary productivity. However, it remains unclear if and how N-induced P limitation varies over time. Soil extracellular phosphatases catalyze the hydrolysis of P from soil organic matter, an important adaptive mechanism for ecosystems to cope with N-induced P limitation. Here we show, using a meta-analysis of 140 studies and 668 observations worldwide, that N stimulation of soil phosphatase activity diminishes over time. Whereas short-term N loading (≤5 years) significantly increased soil phosphatase activity by 28%, long-term N loading had no significant effect. Nitrogen loading did not affect soil available P and total P content in either short- or long-term studies. Together, these results suggest that N-induced P limitation in ecosystems is alleviated in the long-term through the initial stimulation of soil phosphatase activity, thereby securing P supply to support plant growth. Our results suggest that increases in terrestrial carbon uptake due to ongoing anthropogenic N loading may be greater than previously thought.</span>
Cheng R, Magney TS, Dutta D, Bowling DR, Logan BA, Burns SP, Blanken PD, Grossmann K, Lopez S, Richardson AD, Stutz J, Frankenberg K (2020) Decomposing reflectance spectra to track gross primary production in a subalpine evergreen forest. Biogeosciences 17(18): 4523-4544.
Read PublicationChuckran PF, Huntemann M, Clum A, Foster B, Foster B, Roux S, Palaniappan K, Varghese N, Mukherjee S, Reddy TBK, Daum C, Copeland A, Ivanova NN, Kyrpides NC, Del Rio TG, Eloe-Fadrosh EA, Morrissey EM, Schwartz E, Fofanov V, Hungate B, Dijkstra P (2020) Metagenomes and Metatranscriptomes of a Glucose-Amended Agricultural Soil. Microbiology Resource Announcements 9(44): e00895-20.
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Read Publication<span>The addition of glucose to soil has long been used to study the metabolic activity of microbes in soil; however, the response of the microbial ecophysiology remains poorly characterized. To address this, we sequenced the metagenomes and metatranscriptomes of glucose-amended soil microbial communities in a laboratory incubation.</span>
Cui E, Bian C, Luo Y, Niu S, Wang Y, and Xia J (2020) Spatial variations in terrestrial net ecosystem productivity and its local indicators. Biogeosciences 17: 6237-6246.
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Read Publication<span>Multiple lines of evidence have demonstrated the persistence of global land carbon (C) sink during the past several decades. However, both annual net ecosystem productivity (NEP) and its inter-annual variation (IAV</span><span class="inline-formula"><sub>NEP</sub></span><span>) keep varying over space. Thus, identifying local indicators for the spatially varying NEP and IAV</span><span class="inline-formula"><sub>NEP</sub></span><span> is critical for locating the major and sustainable C sinks on land. Here, based on daily NEP observations from FLUXNET sites and large-scale estimates from an atmospheric-inversion product, we found a robust logarithmic correlation between annual NEP and seasonal carbon uptake–release ratio (i.e. </span><span class="inline-formula"><i>U</i> ∕ <i>R</i></span><span>). The cross-site variation in mean annual NEP could be logarithmically indicated by </span><span class="inline-formula"><i>U</i> ∕ <i>R</i></span><span>, while the spatial distribution of IAV</span><span class="inline-formula"><sub>NEP</sub></span><span> was associated with the slope (i.e. </span><span class="inline-formula"><i>β</i></span><span>) of the logarithmic correlation between annual NEP and </span><span class="inline-formula"><i>U</i> ∕ <i>R</i></span><span>. Among biomes, for example, forests and croplands had the largest </span><span class="inline-formula"><i>U</i> ∕ <i>R</i></span><span> ratio (1.06 </span><span class="inline-formula">±</span><span> 0.83) and </span><span class="inline-formula"><i>β</i></span><span> (473 </span><span class="inline-formula">±</span><span> 112 g C m</span><span class="inline-formula"><sup>−2</sup></span><span> yr</span><span class="inline-formula"><sup>−1</sup></span><span>), indicating the highest NEP and IAV</span><span class="inline-formula"><sub>NEP</sub></span><span> in forests and croplands, respectively. We further showed that these two simple indicators could directly infer the spatial variations in NEP and IAV</span><span class="inline-formula"><sub>NEP</sub></span><span> in global gridded NEP products. Overall, this study provides two simple local indicators for the intricate spatial variations in the strength and stability of land C sinks. These indicators could be helpful for locating the persistent terrestrial C sinks and provide valuable constraints for improving the simulation of land–atmospheric C exchanges.</span>
Day NJ, White AL, Johnstone JF, Degré-Timmons G, Cumming SG, Mack MC, Turetsky MR, Walker XJ, Baltzer JL (2020) Fire characteristics and environmental conditions shape plant communities via regeneration strategy. Ecography 43(10): 1464-1474.
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Read Publication<span>Climate change is altering disturbance regimes outside historical norms, which can impact biodiversity by selecting for plants with particular traits. The relative impact of disturbance characteristics on plant traits and community structure may be mediated by environmental gradients. We aimed to understand how wildfire impacted understory plant communities and plant regeneration strategies along gradients of environmental conditions and wildfire characteristics in boreal forests. We established 207 plots (60?m2) in recently burned stands and 133 plots in mature stands with no recent fire history in comparable gradients of stand type, site moisture (drainage) and soil organic layer (SOL) depth in two ecozones in Canada's Northwest Territories. At each plot, we recorded all vascular plant taxa in the understory and measured the regeneration strategy (seeder, resprouter, survivor) in burned plots, along with seedbed conditions (mineral soil and bryophyte cover). Dispersal, longevity and growth form traits were determined for each taxon. Fire characteristics measured included proportion of pre-fire SOL combusted (fire severity), date of burn (fire seasonality) and pre-fire stand age (time following fire). Results showed understory community composition was altered by fire. However, burned and mature stands had similar plant communities in wet sites with deep SOL. In the burned plots, regeneration strategies were determined by fire severity, drainage and pre- and post-fire SOL depth. Resprouters were more common in wet sites with deeper SOL and lower fire severity, while seeders were associated with drier sites with thinner SOL and greater fire severity. This led to drier burned stands being compositionally different from their mature counterparts and seedbed conditions were important. Our study highlights the importance of environment?wildfire interactions in shaping plant regeneration strategies and patterns of understory plant community structure across landscapes, and the overriding importance of SOL depth and site drainage in mediating fire severity, plant regeneration and community structure.</span>
Dieleman CM, Rogers BM, Potter S, Veraverbeke S, Johnstone JF, Laflamme J, Solvik K, Walker XJ, Mack MC, Turetsky MR (2020) Wildfire combustion and carbon stocks in the southern Canadian boreal forest: Implications for a warming world. Global Change Biology 26(11): 60626079.
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Read Publication<span>Boreal wildfires are increasing in intensity, extent, and frequency, potentially intensifying carbon emissions and transitioning the region from a globally significant carbon sink to a source. The productive southern boreal forests of central Canada already experience relatively high frequencies of fire, and as such may serve as an analog of future carbon dynamics for more northern forests. Fire?carbon dynamics in southern boreal systems are relatively understudied, with limited investigation into the drivers of pre-fire carbon stocks or subsequent combustion. As part of NASA's Arctic-Boreal Vulnerability Experiment, we sampled 79 stands (47 burned, 32 unburned) throughout central Saskatchewan to characterize above- and belowground carbon stocks and combustion rates in relation to historical land use, vegetation characteristics, and geophysical attributes. We found southern boreal forests emitted an average of 3.3 ± 1.1 kg C/m2 from field sites. The emissions from southern boreal stands varied as a function of stand age, fire weather conditions, ecozone, and soil moisture class. Sites affected by historical timber harvesting had greater combustion rates due to faster carbon stock recovery rates than sites recovering from wildfire events, indicating that different boreal forest land use practices can generate divergent carbon legacy effects. We estimate the 2015 fire season in Saskatchewan emitted a total of 36.3 ± 15.0 Tg C, emphasizing the importance of southern boreal fires for regional carbon budgets. Using the southern boreal as an analog, the northern boreal may undergo fundamental shifts in forest structure and carbon dynamics, becoming dominated by stands <70 years old that hold 2?7 kg C/m2 less than current mature northern boreal stands. Our latitudinal approach reinforces previous studies showing that northern boreal stands are at a high risk of holding less carbon under changing disturbance conditions.</span>
Feng J, Wang C, Lei J, Yang Y, Yan Q, Zhou X, Tao X, Ning D, Yuan MM, Zin Y, Shi ZJ, Guo X, He Z, Van Nostrand JD, Wu L, Bracho-Garillo, Rosvel G, Penton CR, Cole JR, Konstantinidis KT, Luo Y, Schuur EAG, Tiedje JM, Zhou J (2020) Warming-induced permafrost thaw exacerbates tundra soil carbon decomposition mediated by microbial community. Microbiome 8(1): 3.
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Read Publication<span>It is well-known that global warming has effects on high-latitude tundra underlain with permafrost. This leads to a severe concern that decomposition of soil organic carbon (SOC) previously stored in this region, which accounts for about 50% of the world’s SOC storage, will cause positive feedback that accelerates climate warming. We have previously shown that short-term warming (1.5 years) stimulates rapid, microbe-mediated decomposition of tundra soil carbon without affecting the composition of the soil microbial community (based on the depth of 42684 sequence reads of 16S rRNA gene amplicons per 3 g of soil sample).</span>
Guo D, Li X, Wang J, Niu D, Guo W, Fu H, Luo Y (2020) Edaphic and microbial determinants of the residence times of active and slow C pools on the Tibetan Plateau. Geoderma 357: 113942.
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Read Publication<span>One major source of uncertainties in the estimation of soil organic carbon (SOC) dynamics is carbon (C) residence time, an important parameter in terrestrial ecosystem C cycling models. To better predict terrestrial dynamics, C residence time and its controlling factors need to be well quantified and investigated. In this study, we applied a data assimilation approach to quantify the residence times of different soil C pools on the Tibetan Plateau, based on incubated soil carbon dioxide (CO2) efflux data. We also assessed the effects of soil properties on C residence times. Our results showed that the residence times of active (RTA) and slow (RTS) C pools were well estimated through data assimilation. Soil physical, chemical, and microbial properties significantly regulated RTA and RTS. The RTA was higher in soils with high clay contents (R2 = 0.52, p < 0.01) and low microbial quotients (qMB) (R2 = 0.55, p < 0.01), whereas, the RTS was higher in soils with high clay contents (R2 = 0.76, p < 0.01), high C:N ratios (R2 = 0.44, p < 0.05), low qMBvalues (R2 = 0.50, p < 0.05), and low soil pH values (R2 = 0.80, p < 0.01). Structural equation modeling (SEM) analyses indicated that the model could explain 55% and 91% of the variations in the RTA and RTS, respectively. The qMB was the key factor in regulating the RTA, while soil pH was the crucial variable in controlling the RTS. These results demonstrated that the major controlling factors on the RTA and RTS were different. Considering these variables and their different controls on residence times of different soil C pools could provide more accurate estimation of terrestrial C cycles, and prediction of future C-climate feedbacks.</span>
Guo JS, Gear L, Hultine KR, Koch GW, Ogle K (2020) Non-structural carbohydrate dynamics associated with antecedent stem water potential and air temperature in a dominant desert shrub. Plant, Cell & Environment 43(6): 1467-1483.
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Read Publication<span>Non-structural carbohydrates (NSCs) are necessary for plant growth and affected by plant water status, but the temporal dynamics of water stress impacts on NSC are not well understood. We evaluated how seasonal NSC concentrations varied with plant water status (predawn xylem water potential, ) and air temperature (T) in the evergreen desert shrub Larrea tridentata. Aboveground sugar and starch concentrations were measured weekly or monthly for ~1.5 years on 6 -12 shrubs simultaneously instrumented with automated stem psychrometers; leaf photosynthesis (Anet) was measured monthly for 1 year. Leaf sugar increased during the dry, premonsoon period, associated with lower (greater water stress) and high T. Leaf sugar accumulation coincided with declines in leaf starch and stem sugar, suggesting the prioritization of leaf sugar during low photosynthetic uptake. Leaf starch was strongly correlated with Anet and peaked during the spring and monsoon seasons, while stem starch remained relatively constant except for depletion during the monsoon. Recent photosynthate appeared sufficient to support spring growth, while monsoon growth required the remobilization of stem starch reserves. The coordinated responses of different NSC fractions to water status, photosynthesis, and growth demands suggest that NSCs serve multiple functions under extreme environmental conditions, including severe drought.</span>
Guo JS, Hultine KR, Koch GW, Kropp H, Ogle K (2020) Temporal shifts in iso/anisohydry revealed from daily observations of plant water potential in a dominant desert shrub. New Phytologist 225(2): 713-726.
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Read PublicationPlant species are characterized along a spectrum of isohydry to anisohydry depending on their regulation of water potential (?), but the plasticity of hydraulic strategies is largely unknown. The role of environmental drivers was evaluated in the hydraulic behavior of Larrea tridentata, a drought-tolerant desert shrub that withstands a wide range of environmental conditions. With a 1.5 yr time-series of 2324 in situ measurements of daily predawn and midday ?, the temporal variability of hydraulic behavior was explored in relation to soil water supply, atmospheric demand and temperature. Hydraulic behavior in Larrea was highly dynamic, ranging from partial isohydry to extreme anisohydry. Larrea exhibited extreme anisohydry under wet soil conditions corresponding to periods of high productivity, whereas partial isohydry was exhibited after prolonged dry or cold conditions, when productivity was low. Environmental conditions can strongly influence plant hydraulic behavior at relatively fast timescales, which enhances our understanding of plant drought responses. Although species may exhibit a dominant hydraulic behavior, variable environmental conditions can prompt plasticity in ? regulation, particularly for species in seasonally dry climates.
Hartsell JA, Copeland SM, Munson SM, Butterfield BJ, Bradford JB (2020) Gaps and hotspots in the state of knowledge of pinyon-juniper communities. Forest Ecology and Management 455: 117628.
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Read PublicationPinyon-juniper (PJ) plant communities cover a large area across North America and provide critical habitat for wildlife, biodiversity and ecosystem functions, and rich cultural resources. These communities occur across a variety of environmental gradients, disturbance regimes, structural conditions and species compositions, including three species of juniper and two species of pinyon. PJ communities have experienced substantial changes in recent decades and identifying appropriate management strategies for these diverse communities is a growing challenge. Here, we surveyed the literature and compiled 441 studies to characterize patterns in research on PJ communities through time, across geographic space and climatic conditions, and among focal species. We evaluate the state of knowledge for three focal topics: 1) historical stand dynamics and responses to disturbance, 2) land management actions and their effects, and 3) potential future responses to changing climate. We identified large and potentially important gaps in our understanding of pinyon-juniper communities both geographically and topically. The effect of drought on Pinus edulis, the pinyon pine species in eastern PJ communities was frequently addressed, while few studies focused on drought effects on Pinus monophylla, which occurs in western PJ communities. The largest proportion of studies that examined land management actions only measured their effects for one year. Grazing was a common land-use across the geographic range of PJ communities yet was rarely studied. We found only 39 studies that had information on the impacts of anthropogenic climate change and most were concentrated on Pinus edulis. These results provide a synthetic perspective on PJ communities that can help natural resource managers identify relevant knowledge needed for decision-making and researchers design new studies to fill important knowledge gaps.
Haubensak KA, Grove S, Foster J, Parker IM (2020) Chemical and mechanical control of the invasive shrub Cytisus scoparius in forest clearings in western Washington, USA. Invasive Plant Science and Management 13(1): 30-36.
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Read Publication<span>We conducted a large-scale, multiple-year study in harvested areas of Douglas-fir (Pseudotsuga menziesii [Mirbel] Franco) forests in western Washington, examining the effectiveness of control methods on the widespread invasive shrub Scotch broom [Cytisus scoparius (L.) Link]. We tested both chemical and physical control methods, using three different approaches that are management relevant: (1) triclopyr, a POST herbicide, at different times of year and on different-sized plants; (2) cutting (or brushcutting) of mature individuals; and (3) scarification of soil surface to remove seedlings once versus multiple times. We measured initial mortality, seed germination, and percent cover of C. scoparius in plots for 3 yr following treatments. Triclopyr treatment resulted in greater mortality and reduced percent cover compared with all other treatments with the effect persisting for 2 yr after spraying. Further, triclopyr had the same effect on C. scoparius cover and mortality irrespective of time of year applied. Similar to soil scarification, triclopyr treatments resulted in a flush of seedlings, suggesting that removal of conspecific competitors and not soil disturbance per se promotes seed germination. Brushcutting was generally effective in reducing C. scoparius cover in the short term, but effects did not persist as long as triclopyr treatments, in part due to large differences in stump resprouting rates across sites. Soil scarification to remove seedlings, even over multiple years, did not result in reduced C. scoparius cover. Triclopyr is an effective approach for controlling both emerging and established stands of C. scoparius.</span>
He X, Hou E, Veen GF, Ellwood MDF, Dijkstra P, Sui X, Zhang S, Wen D, Chu C (2020) Soil microbial biomass increases along elevational gradients in the tropics and subtropics but not elsewhere. Global Ecology and Biogeography 29(2): 345-354.
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Read Publication<span>Our aim is to use elevational gradients to quantify the relationship between temperature and ecosystem functioning. Ecosystem functions such as decomposition, nutrient cycling and carbon storage are linked with the amount of microbial biomass in the soil. Previous studies have shown variable relationships between elevation and soil microbial biomass (SMB). Understanding the biological mechanisms linking SMB with elevational gradients will shed light on the environmental impacts of global warming. Location Global. Time period 2002?2018. Major taxa studied Soil microbes. Method We performed a global meta-analysis of the relationships between SMB and elevation. Data were collected from 59 studies of 73 elevational transects from around the world. Results We found no consistent global relationship between SMB and elevation. SMB increased significantly with elevation in the tropics and subtropics, but not in other climate zones. However, we found consistent positive relationships between SMB, soil organic carbon and total nitrogen concentrations. Main conclusions Our results suggest that global warming will impact tropical and subtropical ecosystems more severely than colder regions. Tropical ecosystems, already at risk from species extinctions, will likely experience declines in SMB as the climate warms, resulting in losses of fundamental ecosystem functions such as nutrient cycling and carbon storage.</span>
Hewitt RE, DeVan MR, Lagutina IV, Genet H, McGuire AD, Taylor DL, Mack MC (2020) Mycobiont contribution to tundra plant acquisition of permafrost-derived nitrogen. New Phytologist 226(1): 126-141.
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Read Publication<span>As Arctic soils warm, thawed permafrost releases nitrogen (N) that could stimulate plant productivity and thus offset soil carbon losses from tundra ecosystems. Although mycorrhizal fungi could facilitate plant access to permafrost-derived N, their exploration capacity beyond host plant root systems into deep, cold active layer soils adjacent to the permafrost table is unknown. We characterized root-associated fungi (RAF) that colonized ericoid (ERM) and ectomycorrhizal (ECM) shrub roots and occurred below the maximum rooting depth in permafrost thaw-front soil in tussock and shrub tundra communities. We explored the relationships between root and thaw front fungal composition and plant uptake of a 15N tracer applied at the permafrost boundary. We show that ERM and ECM shrubs associate with RAF at the thaw front providing evidence for potential mycelial connectivity between roots and the permafrost boundary. Among shrubs and tundra communities, RAF connectivity to the thaw boundary was ubiquitous. The occurrence of particular RAF in both roots and thaw front soil was positively correlated with 15N recovered in shrub biomass Taxon-specific RAF associations could be a mechanism for the vertical redistribution of deep, permafrost-derived nutrients, which may alleviate N limitation and stimulate productivity in warming tundra.</span>
Hou E, Luo Y, Kuang Y, Chen C, Lu X, Jiang L, Luo X, Wen D (2020) Global meta-analysis shows pervasive phosphorus limitation of aboveground plant production in natural terrestrial ecosystems. Nature Communications 11(1): 637.
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Read Publication<span>Phosphorus (P) limitation of aboveground plant production is usually assumed to occur in tropical regions but rarely elsewhere. Here we report that such P limitation is more widespread and much stronger than previously estimated. In our global meta-analysis, almost half (46.2%) of 652 P-addition field experiments reveal a significant P limitation on aboveground plant production. Globally, P additions increase aboveground plant production by 34.9% in natural terrestrial ecosystems, which is 7.0–15.9% higher than previously suggested. In croplands, by contrast, P additions increase aboveground plant production by only 13.9%, probably because of historical fertilizations. The magnitude of P limitation also differs among climate zones and regions, and is driven by climate, ecosystem properties, and fertilization regimes. In addition to confirming that P limitation is widespread in tropical regions, our study demonstrates that P limitation often occurs in other regions. This suggests that previous studies have underestimated the importance of altered P supply on aboveground plant production in natural terrestrial ecosystems.</span>
Hou E, Luo Y, Kuang Y, Chen C, Lu X, Jiang L, Luo X, Wen D (2020) Global meta-analysis shows pervasive phosphorus limitation of aboveground plant production in natural terrestrial ecosystems. Nature Communications 11 (637).
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Read PublicationPlants are thought to be limited by phosphorus (P) especially in tropical regions. Here, Hou et al. report a meta-analysis of P fertilization experiments to show widespread P limitation on plant growth across terrestrial ecosystems modulated by climate, ecosystem properties, and fertilization regimes
Jean M, Holland-Moritz H, Melvin AM, Johnstone JF, Mack MC (2020) Experimental assessment of tree canopy and leaf litter controls on the microbiome and nitrogen fixation rates of two boreal mosses. New Phytologist.
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Read Publication<span>Nitrogen (N2)-fixing moss microbial communities play key roles in nitrogen cycling of boreal forests. Forest type and leaf litter inputs regulate moss abundance, but how they control moss microbiomes and N2-fixation remains understudied. We examined the impacts of forest type and broadleaf litter on microbial community composition and N2-fixation rates of Hylocomium splendens and Pleurozium schreberi. We conducted a moss transplant and leaf litter manipulation experiment at three sites with paired paper birch (Betula neoalaskana) and black spruce (Picea mariana) stands in Alaska. We characterized bacterial communities using marker gene sequencing, determined N2-fixation rates using stable isotopes (15N2) and measured environmental covariates. Mosses native to and transplanted into spruce stands supported generally higher N2-fixation and distinct microbial communities compared to similar treatments in birch stands. High leaf litter inputs shifted microbial community composition for both moss species and reduced N2-fixation rates for H. splendens, which had the highest rates. N2-fixation was positively associated with several bacterial taxa, including cyanobacteria. The moss microbiome and environmental conditions controlled N2-fixation at the stand and transplant scales. Predicted shifts from spruce- to deciduous-dominated stands will interact with the relative abundances of mosses supporting different microbiomes and N2-fixation rates, which could affect stand-level N inputs.</span>
Liu XJA, Finley BK, Mau RL, Schwartz E, Dijkstra P, Bowker MA, Hungate BA (2020) The soil priming effect: Consistent across ecosystems, elusive mechanisms. Soil Biology and Biochemistry 140: 107617.
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Read Publication<span>Organic matter input to soils can accelerate the decomposition of native soil carbon (C), a process called the priming effect. Priming is ubiquitous and exhibits some consistent patterns, but a general explanation remains elusive, in part because of variation in the response across different ecosystems, and because of a diversity of proposed mechanisms, including microbial activation, stoichiometry, and community shifts. Here, we conducted five-week incubations of four soils (grassland, piñon-juniper, ponderosa pine, mixed conifer), varying the amount of substrate added (as 13C-glucose, either 350 or 1000 μg C g−1 week−1) and either with no added nitrogen (N), or with sufficient N (as NH4NO3) to bring the C-to-N ratio of the added substrate to 10. Using four different ecosystems enabled testing the generality of mechanisms underlying the priming effect. The responses of priming to the amount and C-to-N ratio of the added substrate were consistent across ecosystems: priming increased with the rate of substrate addition and declined when the C-to-N ratio of the substrate was reduced. However, structural equation models failed to confirm intermediate responses postulated to mediate the priming effect, including responses postulated to be mediated by stoichiometry and microbial activation. Specifically, priming was not clearly associated with changes in microbial biomass or turnover, nor with extracellular enzyme activities or the microbial C-to-N ratio. The strongest explanatory pathways in the structural equation models were the substrate, soil, and C-to-N ratio treatments themselves, with no intermediates, suggesting that either these measurements lacked sufficient sensitivity to reveal causal relationships, or the actual drivers for priming were not included in the ancillary measurements. While we observed consistent changes in priming caused by the amount and C-to-N ratio of the added substrate across a wide array of soils, our findings did not clearly conform to common models offered for the priming effect. Because priming is a residual flux involving diverse substrates of varying chemical composition, a simple and generalizable explanation of the phenomenon may be elusive.</span>
Luo X, Hou E, Chen J, Li J, Zhang L, Zang X, Wen D (2020) Dynamics of carbon, nitrogen, and phosphorus stocks and stoichiometry resulting from conversion of primary broadleaf forest to plantation and secondary forest in subtropical China. CATENA 193: 104606.
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Read Publication<span>Large-scale primary native broadleaf forests (BF) have been converted to secondary forests (SF) and plantation forests (PF) in subtropical China over the past decades. However, how and what magnitude of plant and soil carbon (C), nitrogen (N), and phosphorus (P) stocks and stoichiometry are affected by forest conversion is still vague. Here, we addressed this issue by systematically measuring tree biomass and the C, N, and P concentrations in tree organs and soils (0–100 cm) collected from 300 plots in Fujian province. With forest conversion of BF to PF, the tree C, N, and P stocks declined by 43.8, 47.9, and 63.1%, respectively, and the soil C and N stocks across whole soil depth decreased by 19.1% and 13.0%, respectively, and these decreases were more evident after conversion of BF to PF than SF. However, soil P stock showed a tendency of decreasing at 0–20 cm soil depth but increasing at 20–100 cm soil depth following conversion of BF to SF and PF. This unconformity of the vertical pattern of P stock in contrast to C and N stocks, was perhaps due to higher C and N inputs and greater P uptake from the subsoil and its redistribution to the topsoil in BF than in SF and PF. The tree and soil C, N, and P stoichiometry was strongly related to tree biomass, indicating that tree biomass was a vital factor driving soil inputs and retention of nutrients, and thus affecting their stoichiometry. The leaf N:P ratios ranging from 16.7 to 17.2 at our study sites suggested that co-limitations of N and P for forest growth could occur in the studied region. Our results provided insights into the C, N, and P linkages between soils and trees as affected by forest conversion, and advised that predicting these linkages could be an effective approach to identify the impacts of forest conversion, thereby implementing targeted conservation and rehabilitation actions.</span>
Luo Y, Xia J (2020) A dynamic disequilibrium hypothesis for terrestrial carbon cycle. Biodiversity Science 28(11):1405-1416.
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Read Publication<span>The dynamic equilibrium of mass and energy movement in ecosystems is an important basis for the Earth system to nurture and maintain biodiversity. Since the Industrial Revolution, human activities have caused the carbon exchange between terrestrial ecosystems and the atmosphere to be at dynamic disequilibrium. This paper examines a dynamic disequilibrium hypothesis for the carbon cycle of terrestrial ecosystems. The hypothesis suggests that the dynamic disequilibrium is caused by interactions of four basic properties of internal processes of the terrestrial carbon cycle with five types of external drivers. Based on these internal properties and external drivers, this paper summarizes the expression phenomena of the dynamic disequilibrium of terrestrial carbon cycle at different time and space scales, and discusses its detection methods from the perspective of observations, experiments and models. The dynamic disequilibrium hypothesis for terrestrial carbon cycle not only helps us understand the complex terrestrial carbon-cycle phenomenon, but also provides a new theoretical framework for predicting the future terrestrial carbon sink dynamics.</span>
Meng L, Mao J, Zhou Y, Richardson AD, Lee X, Thornton PE, Ricciuto DM, Li X, Dai Y, Shi X, Jia G (2020) Urban warming advances spring phenology but reduces the response of phenology to temperature in the conterminous United States. Proceedings of the National Academy of Sciences 117(8): 4228.
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Read Publication<span>Cities and their associated urban heat islands are ideal natural laboratories for evaluating the response of plant phenology to warming conditions. In this study, we demonstrate that the satellite-derived start of season for plants occurred earlier but showed less covariation with temperature in most of the large 85 cities across the conterminous United States for the period 2001–2014. The results show a reduction in the response of urban phenology to temperature and imply that, in nonurban environments, the onset of spring phenology will likely advance but will slow down as the general trend toward warming continues.Urbanization has caused environmental changes, such as urban heat islands (UHIs), that affect terrestrial ecosystems. However, how and to what extent urbanization affects plant phenology remains relatively unexplored. Here, we investigated the changes in the satellite-derived start of season (SOS) and the covariation between SOS and temperature (RT) in 85 large cities across the conterminous United States for the period 2001–2014. We found that 1) the SOS came significantly earlier (6.1 ± 6.3 d) in 74 cities and RT was significantly weaker (0.03 ± 0.07) in 43 cities when compared with their surrounding rural areas (P &amp;lt; 0.05); 2) the decreased magnitude in RT mainly occurred in cities in relatively cold regions with an annual mean temperature &amp;lt;17.3 °C (e.g., Minnesota, Michigan, and Pennsylvania); and 3) the magnitude of urban−rural difference in both SOS and RT was primarily correlated with the intensity of UHI. Simulations of two phenology models further suggested that more and faster heat accumulation contributed to the earlier SOS, while a decrease in required chilling led to a decline in RT magnitude in urban areas. These findings provide observational evidence of a reduced covariation between temperature and SOS in major US cities, implying the response of spring phenology to warming conditions in nonurban environments may decline in the warming future.</span>
Papp K, Hungate BA, Schwartz E (2020) Glucose triggers strong taxon-specific responses in microbial growth and activity: insights from DNA and RNA qSIP. Ecology 101(1): e02887.
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Read Publication<span>Growth of soil microorganisms is often described as carbon limited, and adding labile carbon to soil often results in a transient and large increase in respiration. In contrast, soil microbial biomass changes little, suggesting that growth and respiration are decoupled in response to a carbon pulse. Alternatively, measuring bulk responses of the entire community (total respiration and biomass) could mask ecologically important variation among taxa in response to the added carbon. Here, we assessed taxon-specific variation in cellular growth (measured as DNA synthesis) and metabolic activity (measured as rRNA synthesis) following glucose addition to soil using quantitative stable isotope probing with H218O. We found that glucose addition altered rates of DNA and rRNA synthesis, but the effects were strongly taxon specific: glucose stimulated growth and rRNA transcription for some taxa, and suppressed these for others. These contrasting taxon-specific responses could explain the small and transient changes in total soil microbial biomass. Responses to glucose were not well predicted by a priori assignments of taxa into copiotrophic or oligotrophic categories. Across all taxa, rates of DNA and rRNA synthesis changed in parallel, indicating that growth and activity were coupled, and the degree of coupling was unaffected by glucose addition. This pattern argues against the idea that labile carbon addition causes a large reduction in metabolic growth efficiency; rather, the large pulse of respiration observed with labile substrate addition is more likely to be the result of rapid turnover of microbial biomass, possibly due to trophic interactions. Our results support a strong connection between rRNA synthesis and bacterial growth, and indicate that taxon-specific responses among soil bacteria can buffer responses at the scale of the whole community.</span>
Pappalardo P, Ogle K, Hamman EA, Bence JR, Hungate BA (2020) Comparing traditional and Bayesian approaches to ecological meta-analysis. Methods in Ecology and Evolution.
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Read Publication<span>Despite the wide application of meta-analysis in ecology, some of the traditional methods used for meta-analysis may not perform well given the type of data characteristic of ecological meta-analyses. We reviewed published meta-analyses on the ecological impacts of global climate change, evaluating the number of replicates used in the primary studies (ni) and the number of studies or records (k) that were aggregated to calculate a mean effect size. We used the results of the review in a simulation experiment to assess the performance of conventional frequentist and Bayesian meta-analysis methods for estimating a mean effect size and its uncertainty interval. Our literature review showed that ni and k were highly variable, distributions were right-skewed and were generally small (median ni = 5, median k = 44). Our simulations show that the choice of method for calculating uncertainty intervals was critical for obtaining appropriate coverage (close to the nominal value of 0.95). When k was low (<40), 95% coverage was achieved by a confidence interval (CI) based on the t distribution that uses an adjusted standard error (the Hartung?Knapp?Sidik?Jonkman, HKSJ), or by a Bayesian credible interval, whereas bootstrap or z distribution CIs had lower coverage. Despite the importance of the method to calculate the uncertainty interval, 39% of the meta-analyses reviewed did not report the method used, and of the 61% that did, 94% used a potentially problematic method, which may be a consequence of software defaults. In general, for a simple random-effects meta-analysis, the performance of the best frequentist and Bayesian methods was similar for the same combinations of factors (k and mean replication), though the Bayesian approach had higher than nominal (>95%) coverage for the mean effect when k was very low (k < 15). Our literature review suggests that many meta-analyses that used z distribution or bootstrapping CIs may have overestimated the statistical significance of their results when the number of studies was low; more appropriate methods need to be adopted in ecological meta-analyses.</span>
Pellegrini AFA, McLauchlan KK, Hobbie SE, Mack MC, Marcotte AL, Nelson DM, Perakis SS, Reich PB, Whittinghill K (2020) Frequent burning causes large losses of carbon from deep soil layers in a temperate savanna. Journal of Ecology 108(4): 1426-1441.
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Read Publication<span>Fire activity is changing dramatically across the globe, with uncertain effects on ecosystem processes, especially below-ground. Fire-driven losses of soil carbon (C) are often assumed to occur primarily in the upper soil layers because the repeated combustion of above-ground biomass limits organic matter inputs into surface soil. However, C losses from deeper soil may occur if frequent burning reduces root biomass inputs of C into deep soil layers or stimulates losses of C via leaching and priming. To assess the effects of fire on soil C, we sampled 12 plots in a 51-year-long fire frequency manipulation experiment in a temperate oak savanna, where variation in prescribed burning frequency has created a gradient in vegetation structure from closed-canopy forest in unburned plots to open-canopy savanna in frequently burned plots. Soil C stocks were nonlinearly related to fire frequency, with soil C peaking in savanna plots burned at an intermediate fire frequency and declining in the most frequently burned plots. Losses from deep soil pools were significant, with the absolute difference between intermediately burned plots versus most frequently burned plots more than doubling when the full 1 m sample was considered rather than the top 0?20 cm alone (losses of 98.5 Mg C/ha [?76%] and 42.3 Mg C/ha [?68%] in the full 1 m and 0?20 cm layers respectively). Compared to unburned forested plots, the most frequently burned plots had 65.8 Mg C/ha (?58%) less C in the full 1 m sample. Root biomass below the top 20 cm also declined by 39% with more frequent burning. Concurrent fire-driven losses of nitrogen and gains in calcium and phosphorus suggest that burning may increase nitrogen limitation and play a key role in the calcium and phosphorus cycles in temperate savannas. Synthesis. Our results illustrate that fire-driven losses in soil C and root biomass in deep soil layers may be critical factors regulating the net effect of shifting fire regimes on ecosystem C in forest-savanna transitions. Projected changes in soil C with shifting fire frequencies in savannas may be 50% too low if they only consider changes in the topsoil.</span>
Peltier DMP, Guo J, Nguyen P, Bangs M, Gear L, Wilson M, Jeffers S, Samuels-Crow K, Yocom LL, Liu Y, Fell MK, Auty D, Schwalm C, Anderegg WRL, Koch GW, Litvak ME, Ogle K (2020) Temporal controls on crown nonstructural carbohydrates in southwestern US tree species. Tree Physiology.
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Read Publication<span>In trees, large uncertainties remain in how nonstructural carbohydrates (NSCs) respond to variation in water availability in natural, intact ecosystems. Variation in NSC pools reflects temporal fluctuations in supply and demand, as well as physiological coordination across tree organs in ways that differ across species and NSC fractions (e.g., soluble sugars vs starch). Using landscape-scale crown (leaves and twigs) NSC concentration measurements in three foundation tree species (Populus tremuloides, Pinus edulis, Juniperus osteosperma), we evaluated in situ, seasonal variation in NSC responses to moisture stress on three timescales: short-term (via predawn water potential), seasonal (via leaf δ13C) and annual (via current year’s ring width index). Crown NSC responses to moisture stress appeared to depend on hydraulic strategy, where J. osteosperma appears to regulate osmotic potentials (via higher sugar concentrations), P. edulis NSC responses suggest respiratory depletion and P. tremuloides responses were consistent with direct sink limitations. We also show that overly simplistic models can mask seasonal and tissue variation in NSC responses, as well as strong interactions among moisture stress at different timescales. In general, our results suggest large seasonal variation in crown NSC concentrations reflecting the multiple cofunctions of NSCs in plant tissues, including storage, growth and osmotic regulation of hydraulically vulnerable leaves. We emphasize that crown NSC pool size cannot be viewed as a simple physiological metric of stress; in situ NSC dynamics are complex, varying temporally, across species, among NSC fractions and among tissue types.</span>
Peng W, Song T, Du H, Chen H, Zeng F, Liu Y, Luo Y, Tan W (2020) Inconsistent diversity patterns of soil fungi and woody plants among habitat types in a karst broadleaf forest. Forest Ecology and Management 474: 118367.
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Read Publication<span>The diversity patterns of macroorganisms (i.e., plants) among different habitats have been well documented, however, those of microorganisms (i.e., fungi) as well as the relationships between them are still unclear. Here, we tested whether and to what degree fungal diversity was related to habitat types and compared diversity patterns of woody plants and soil fungi. We carried out field investigations on soil fungi in different habitat types (i.e., valleys, foothills, hillsides, and hilltops) in a 25-ha karst broadleaf forest in Southwest China. The tree richness, Shannon index, and Simpson index significantly increased from valleys to hilltops. While the soil fungal N1 diversity (the exponential Shannon index) marginally increased toward valleys, fungal N0 (richness) and N2 (the inverse Simpson index) diversity exhibited significantly reduced and increased patterns, respectively, from valleys to hilltops. The major fungal functional groups (i.e., EcM, AM, saprotrophic, and pathogenic fungi) showed similar increasing richness patterns in valleys. Moreover, woody plant alpha diversity was an important indicator of fungal functional groups except for EcM and AM fungi. In addition, woody plants increased in species turnover rate (βSIM) from valleys to hilltops, while fungal species had a concave distribution. The patterns of nestedness (βSNE) for tree species decreased from valleys to hilltops, while the opposite was true for soil fungal species. Our findings indicated that the diversity patterns of woody plants and fungi were inconsistent among habitat types, and the relationships between fungal and woody plant communities depended on habitat types in the karst forest.</span>
Pugh TAM, Rademacher TT, Shafer SL, Steinkamp J, Barichivich J, Leckage B, Havard V, Harper A, Heinke J, Nishina K, Rammig A, Sato H, Arneth A, Hantson S, Hickler T, Kautz M, Quesada B, Smith B, Thonicke K (2020) Understanding the uncertainty in global forest carbon turnover. Biogeosciences Discuss 2020: 1-44.
Read PublicationQian H, Huang S, Chen J, Wang L, Hungate BA, Van Kessel C, Zhang J, Deng A, Jiang Y, Van Groenigen KJ, Zhang W (2020) Lower-than-expected CH4 emissions from rice paddies with rising CO2 concentrations. Global Change Biology 26(4): 2368-2376.
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Read Publication<span>Elevated atmospheric CO2 (eCO2) generally increases carbon input in rice paddy soils and stimulates the growth of methane-producing microorganisms. Therefore, eCO2 is widely expected to increase methane (CH4) emissions from rice agriculture, a major source of anthropogenic CH4. Agricultural practices strongly affect CH4 emissions from rice paddies as well, but whether these practices modulate effects of eCO2 is unclear. Here we show, by combining a series of experiments and meta-analyses, that whereas eCO2 strongly increased CH4 emissions from paddies without straw incorporation, it tended to reduce CH4 emissions from paddy soils with straw incorporation. Our experiments also identified the microbial processes underlying these results: eCO2 increased methane-consuming microorganisms more strongly in soils with straw incorporation than in soils without straw, with the opposite pattern for methane-producing microorganisms. Accounting for the interaction between CO2 and straw management, we estimate that eCO2 increases global CH4 emissions from rice paddies by 3.7%, an order of magnitude lower than previous estimates. Our results suggest that the effect of eCO2 on CH4 emissions from rice paddies is smaller than previously thought and underline the need for judicious agricultural management to curb future CH4 emissions.</span>
Qiu T, Song C, Clark JS, Seyednasrollah B, Rathnayaka B, Li J (2020) Understanding the continuous phenological development at daily time step with a Bayesian hierarchical space-time model: impacts of climate change and extreme weather events. Remote Sensing of Environment 247: 111956.
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Read Publication<span>The impacts of climate change and extreme weather events (e.g. frost-, heat-, drought-, and heavy rainfall events) on the continuous phenological development over the entire seasonal cycle remained poorly understood. Previous studies mainly focused on modeling key phenological transition dates (e.g. discrete timing of spring bud-break and fall senescence) based on aggregated climate variables (e.g. mean temperature, growing-degree days). We developed and evaluated a Bayesian Hierarchical Space-Time model for Land Surface Phenology (BHST-LSP) to synthesize remotely sensed vegetation greenness with climate covariates at a daily temporal scale from 1981 to 2014 across the entire conterminous United States. The BHST-LSP model incorporated both temporal and spatial information and exhibited high predictive power in simulating daily phenological development with an overall out-of-sample R2 of 0.80 ± 0.17 and 0.72 ± 0.20 for spring and fall phenology, respectively. The overall out-of-sample normalized root mean square errors were 9.3% ± 6.1% and 9.9% ± 5.2% between the observed and predicted vegetation greenness for spring and fall phenology, respectively. We found that a fast increase of temperature can accelerate the speed of spring green-up while a slow decrease of temperature can lead to a decelerated fall brown-down. Increasing accumulated precipitation can benefit daily phenological development over an entire growing season, while extreme rainfall events can have the opposite effects. More frequent frost events could slow spring leaf expansion and accelerate fall leaf senescence. Impacts of extreme heat events were complex and depended on water availability. Cropland in the Midwest as well as evergreen needleleaf forest along the coastal regions showed relatively strong resistance to drought events compared to other land cover types. The BHST-LSP model can be used to forecast vegetation phenology given future climate projection, thus providing valuable information for adopting climate change adaptation and mitigation measures.</span>
Rodenhizer H, Ledman J, Mauritz M, Natali SM, Pegoraro E, Plaza C, Romano E, Schädel C, Taylor M, Schuur E (2020) Carbon thaw rate doubles when accounting for subsidence in a permafrost warming experiment. Journal of Geophysical Research: Biogeosciences 125(6): e2019JG005528.
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Read Publication<span>Permafrost thaw is typically measured with active layer thickness, or the maximum seasonal thaw measured from the ground surface. However, previous work has shown that this measurement alone fails to account for ground subsidence and therefore underestimates permafrost thaw. To determine the impact of subsidence on observed permafrost thaw and thawed soil carbon stocks, we quantified subsidence using high-accuracy GPS and identified its environmental drivers in a permafrost warming experiment near the southern limit of permafrost in Alaska. With permafrost temperatures near 0°C, 10.8 cm of subsidence was observed in control plots over 9 years. Experimental air and soil warming increased subsidence by five times and created inundated microsites. Across treatments, ice and soil loss drove 85?91% and 9?15% of subsidence, respectively. Accounting for subsidence, permafrost thawed between 19% (control) and 49% (warming) deeper than active layer thickness indicated, and the amount of newly thawed carbon within the active layer was between 37% (control) and 113% (warming) greater. As additional carbon thaws as the active layer deepens, carbon fluxes to the atmosphere and lateral transport of carbon in groundwater could increase. The magnitude of this impact is uncertain at the landscape scale, though, due to limited subsidence measurements. Therefore, to determine the full extent of permafrost thaw across the circumpolar region and its feedback on the carbon cycle, it is necessary to quantify subsidence more broadly across the circumpolar region.</span>
Rubin RL, Jones AN, Hayer M, Shuman-Goodier ME, Andrews LV, Hungate BA (2020) Opposing effects of bacterial endophytes on biomass allocation of a wild donor and agricultural recipient. FEMS Microbiology Ecology 96(3).
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Read Publication<span>Root endophytes are a promising tool for increasing plant growth, but it is unclear whether they perform consistently across plant hosts. We characterized the blue grama (Bouteloua gracilis) root microbiome using two sequencing methods, quantified the effects of root endophytes in the original host (blue grama) and an agricultural recipient, corn (Zea mays), under drought and well-watered conditions and examined in vitro mechanisms for plant growth promotion. 16S rRNA amplicon sequencing revealed that the blue grama root microbiome was similar across an elevation gradient, with the exception of four genera. Culturing and Sanger sequencing revealed eight unique endophytes belonging to the genera Bacillus, Lysinibacillus and Pseudomonas. All eight endophytes colonized corn roots, but had opposing effects on aboveground and belowground biomass in each plant species: they increased blue grama shoot mass by 45% (19) (mean +/− SE) while decreasing corn shoot mass by 10% (19), and increased corn root:shoot by 44% (7), while decreasing blue grama root:shoot by 17% (7). Furthermore, contrary to our expectations, endophytes had stronger effects on plant growth under well-watered conditions rather than drought conditions. Collectively, these results suggest that ecological features, including host identity, bacterial traits, climate conditions and morphological outcomes, should be carefully considered in the design and implementation of agricultural inocula.</span>
Samuels-Crow KE, Ogle K, Litvak ME (2020) Atmosphere-soil Interactions govern ecosystem flux sensitivity to environmental Conditions in semiarid woody ecosystems over varying timescales. Journal of Geophysical Research: Biogeosciences e2019JG005554.
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Read Publication<span>Water and CO2 flux responses (e.g., evapotranspiration [ET] and net ecosystem exchange [NEE]) to environmental conditions can provide insights into how climate change will affect the terrestrial water and carbon budgets, especially in sensitive semiarid ecosystems. Here, we evaluated sensitivity of daily ET and NEE to current and antecedent (past) environment conditions, including atmospheric (vapor pressure deficit [VPD] and air temperature [Tair]) and moisture (precipitation and soil water) drivers. We focused on two common southwestern U.S. (?Southwest?) biomes: pinyon-juniper woodland (Pinus edulis, Juniperus monosperma) and ponderosa pine forest (Pinus ponderosa). Due to differences in aridity, rooting patterns, and plant physiological strategies (stomatal and hydraulic traits), we expected ET and NEE in these ecosystems to respond differently to atmospheric and moisture drivers, with longer response timescales in the drier pinyon-juniper woodland. Net sensitivity to drivers varied temporally in both ecosystems, reflecting the integrated influence of interacting drivers and antecedent precipitation patterns. NEE sensitivity to VPD and soil moisture (and ET sensitivity to deep soil moisture [Sdeep]) was higher in the ponderosa forest. ET and NEE in both ecosystems responded almost instantaneously to Tair, VPD, and shallow soil moisture (Sshall), and increases in any of these drivers weakened the carbon sink and enhanced water loss. Conversely, Sdeep and precipitation influenced ET and NEE over longer timescales (days to months, respectively), and higher Sdeep enhanced the carbon sink. As climate changes, these results suggest hotter and drier conditions will weaken the carbon sink and exacerbate water loss from Southwest pinyon-juniper and ponderosa ecosystems.</span>
Schaedel C, Beem-Miller J, Aziz Rad M, Crow SE, Hicks Pries CE, Ernakovich J, Hoyt AM, Plante A, Stoner S, Treat CC, Sierra CA (2020) Decomposability of soil organic matter over time: the Soil Incubation Database (SIDb, version 1.0) and guidance for incubation procedures. Earth Systems Science Data 12:1511-1524.
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Read Publication<span>The magnitude of carbon (C) loss to the atmosphere via microbial decomposition is a function of the amount of C stored in soils, the quality of the organic matter, and physical, chemical, and biological factors that comprise the environment for decomposition. The decomposability of C is commonly assessed by laboratory soil incubation studies that measure greenhouse gases mineralized from soils under controlled conditions. Here, we introduce the Soil Incubation Database (SIDb) version 1.0, a compilation of time series data from incubations, structured into a new, publicly available, open-access database of C flux (carbon dioxide, </span><span class="inline-formula">CO<sub>2</sub></span><span>, or methane, </span><span class="inline-formula">CH<sub>4</sub></span><span>). In addition, the SIDb project also provides a platform for the development of tools for reading and analysis of incubation data as well as documentation for future use and development. In addition to introducing SIDb, we provide reporting guidance for database entry and the required variables that incubation studies need at minimum to be included in SIDb. A key application of this synthesis effort is to better characterize soil C processes in Earth system models, which will in turn reduce our uncertainty in predicting the response of soil C decomposition to a changing climate. We demonstrate a framework to fit curves to a number of incubation studies from diverse ecosystems, depths, and organic matter content using a built-in model development module that integrates SIDb with the existing SoilR package to estimate soil C pools from time series data. The database will help bridge the gap between point location measurements, which are commonly used in incubation studies, and global remote-sensed data or data products derived from models aimed at assessing global-scale rates of decomposition and C turnover. The SIDb version 1.0 is archived and publicly available at </span><a href="https://doi.org/10.5281/zenodo.3871263">https://doi.org/10.5281/zenodo.3871263</a><span> (Sierra et al., 2020), and the database is managed under a version-controlled system and centrally stored in GitHub (</span><span class="uri"><a href="https://github.com/SoilBGC-Datashare/sidb" target="_blank" rel="noopener noreferrer">https://github.com/SoilBGC-Datashare/sidb</a></span><span>, last access: 26 June 2020).</span>
Schwalm CR, Huntinzger DN, Michalak AM, Schafer K, Fisher JB, Fang Y, Wei Y (2020) Modeling suggests fossil fuel emissions have been driving increased land carbon uptake since the turn of the 20th Century. Scientific Reports 10(1): 9059.
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Read Publication<span>Terrestrial vegetation removes CO2 from the atmosphere; an important climate regulation service that slows global warming. This 119 Pg C per annum transfer of CO2 into plants—gross primary productivity (GPP)—is the largest land carbon flux globally. While understanding past and anticipated future GPP changes is necessary to support carbon management, the factors driving long-term changes in GPP are largely unknown. Here we show that 1901 to 2010 changes in GPP have been dominated by anthropogenic activity. Our dual constraint attribution approach provides three insights into the spatiotemporal patterns of GPP change. First, anthropogenic controls on GPP change have increased from 57% (1901 decade) to 94% (2001 decade) of the vegetated land surface. Second, CO2 fertilization and nitro gen deposition are the most important drivers of change, 19.8 and 11.1 Pg C per annum (2001 decade) respectively, especially in the tropics and industrialized areas since the 1970’s. Third, changes in climate have functioned as fertilization to enhance GPP (1.4 Pg C per annum in the 2001 decade). These findings suggest that, from a land carbon balance perspective, the Anthropocene began over 100 years ago and that global change drivers have allowed GPP uptake to keep pace with anthropogenic emissions.</span>
Seyednasrollah B, Bowling DR, Cheng R, Logan BA, Magney TS, Frankenberg C, Yang JC, Young AM, Hufkens K, Arain MA, Black TA, Blanken PD, Bracho R, Jassal R, Hollinger DY, Law BE, Nesic Z, Richardson AD (2020) Seasonal variation in the canopy color of temperate evergreen conifer forests. New Phytologist.
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Read Publication<span>Evergreen conifer forests are the most prevalent land cover type in North America. Seasonal changes in the color of evergreen forest canopies have been documented with near-surface remote sensing, but the physiological mechanisms underlying these changes, and the implications for photosynthetic uptake, have not been fully elucidated. Here, we integrate on-the-ground phenological observations, leaf-level physiological measurements, near surface hyperspectral remote sensing and digital camera imagery, tower-based CO2 flux measurements, and a predictive model to simulate seasonal canopy color dynamics. We show that seasonal changes in canopy color occur independently of new leaf production, but track changes in chlorophyll fluorescence, the photochemical reflectance index, and leaf pigmentation. We demonstrate that at winter-dormant sites, seasonal changes in canopy color can be used to predict the onset of canopy-level photosynthesis in spring, and its cessation in autumn. Finally, we parameterize a simple temperature-based model to predict the seasonal cycle of canopy greenness, and we show that the model successfully simulates interannual variation in the timing of changes in canopy color. These results provide mechanistic insight into the factors driving seasonal changes in evergreen canopy color and provide opportunities to monitor and model seasonal variation in photosynthetic activity using color-based vegetation indices.</span>
Seyednasrollah B, Clark JS (2020) Where resource-acquisitive species are located: the role of habitat heterogeneity. Geophysical Research Letters 47(8): e2020GL087626.
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Read Publication<span>Rising temperatures with increased drought pose two challenges for management of future biodiversity. First, are the most vulnerable species concentrated in specific regions and habitats? Second, where can landscape heterogeneity potentially mitigate impacts? We conducted a comprehensive trait analysis of forest plots spanning the eastern United States to quantify how resource-acquisitive species respond to moisture-soil-climate interactions. We found that resource-acquisitive species, including nutrient-acquisitive and moisture-acquisitive species, respond disproportionately to environmental gradients, and their response is largely explained by soil variation. We showed that the strong boundary of resource-acquisitive species occurs near the last glacial limit, highlighting one of the clearest indicators of soil controls. Although local soil moisture may reduce drought-induced stress for moisture-acquisitive species, nutrient-acquisitive species remain vulnerable on wet soils in dry climates. The results suggest that theories explaining species distributions should devote close attention to the combination of local drainage and soil type.</span>
Siders AC, Compson ZG, Hungate BA, Dijkstra P, Koch GW, Marks JC (2020) The Influence of Leaf Type on Carbon and Nitrogen Assimilation by Aquatic Invertebrate Communities: A New Perspective on Trophic Efficiency. Ecosystems.
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Read Publication<span>Despite abounding evidence that leaf litter traits can predict decomposition rate, the way these traits influence trophic efficiency and element transfer to higher trophic levels is not resolved. Here, we used litter labeled with 13C and 15N stable isotopes to trace fluxes of litter C and N from four leaf types to freshwater invertebrate communities. We measured absolute (mg C or N) and relative assimilation (percentage of litter C or N incorporated into invertebrate biomass relative to C and N lost during decomposition). Four patterns emerged: (1) Invertebrate communities assimilated more C and N from slowly decomposing litter than communities feeding on rapidly decomposing litter; (2) absolute assimilation of both C and N in leaf packs was positively correlated with the relative biomass of invertebrate taxa in leaf packs; (3) Chironomidae larvae, which colonize packs in the early decomposition stages, assimilated the most C and N by the end of the 35-day experiment; and (4) most taxa, spanning five functional feeding groups (collector–gatherers, shredders, collector–filterers, scrapers, and predators), showed similar patterns in both absolute and relative assimilation across leaf types. These results challenge traditional views of litter quality by demonstrating that trophic efficiency is negatively associated with decomposition rate across these four leaf types.</span>
Sieradzki ET, Koch BJ, Greenlon A, Sachdeva R, Malmstrom RR, Mau RL, Blazewics SJ, Firestone MK, Hofmockel K, Schwartz E, Hungate BA, Pett-Ridge J (2020) Measurement error and resolution in quantitative stable isotope probing: implications for experimental design. bioRxiv 2020.02.25.965764.
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Read Publication<span>Quantitative stable isotope probing (qSIP) estimates the degree of incorporation of an isotope tracer into nucleic acids of metabolically active organisms and can be applied to microorganisms growing in complex communities, such as the microbiomes of soil or water. As such, qSIP has the potential to link microbial biodiversity and biogeochemistry. As with any technique involving quantitative estimation, qSIP involves measurement error; a more complete understanding of error, precision and statistical power will aid in the design of qSIP experiments and interpretation of qSIP data. We used several existing qSIP datasets of microbial communities found in soil and water to evaluate how variance in the estimate of isotope incorporation depends on organism abundance and on the resolution of the density fractionation scheme. We also assessed statistical power for replicated qSIP studies, and sensitivity and specificity for unreplicated designs. We found that variance declines as taxon abundance increases. Increasing the number of density fractions reduces variance, although the benefit of added fractions declines as the number of fractions increases. Specifically, nine fractions appear to be a reasonable tradeoff between cost and precision for most qSIP applications. Increasing replication improves power and reduces the minimum detectable threshold for inferring isotope uptake to 5 atom%. Finally, we provide evidence for the importance of internal standards to calibrate the %GC to mean weighted density regression per sample. These results should benefit those designing future SIP experiments, and provide a reference for metagenomic SIP applications where financial and computational limitations constrain experimental scope.Importance One of the biggest challenges in microbial ecology is correlating the identity of microorganisms with the roles they fulfill in natural environmental systems. Studies of microbes in pure culture reveal much about genomic content and potential functions, but may not reflect an organism’s activity within its natural community. Culture-independent studies supply a community-wide view of composition and function in the context of community interactions, but fail to link the two. Quantitative stable isotope probing (qSIP) is a method that can link the identity and function of specific microbes within a naturally occurring community. Here we explore how the resolution of density-gradient fractionation affects the error and precision of qSIP results, how they may be improved via additional replication, and cost-benefit balanced scenarios for SIP experimental design.</span>
Silva LVB, Vasconcelos HL, Mack MC, Ferreira AS, Bruna EM (2020) Effects of experimental nitrogen enrichment on soil properties and litter decomposition in a Neotropical savanna. Austral Ecology 45(8):1093-1102.
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Read Publication<span>The amount of reactive nitrogen has more than doubled in terrestrial ecosystems due to human activities such fertiliser application that is predicted to increase dramatically in coming decades. We conducted a 3-year experiment in a Neotropical savanna in which we determined the effects of increased N deposition on litter decomposition in plots subjected to different levels of N addition (50 kg N ha?1 year?2, 20 kg N ha?1 year?2, or no N addition). For this, we compared the litter decomposition from the bunchgrass Tristachya leiostachya using litter collected from plots with different N addition treatments. Five randomly selected bags of litter from each N addition treatment (origin) were distributed to each plot (destination). We also compared litter nitrogen (N) concentration and indicators of microbial activity (basal respiration, carbon of microbial biomass, metabolic quotient, enzyme activity) in all plots. We found that nitrogen addition influences litter decay, but in idiosyncratic ways that differ between years. In year 1, litter decomposed faster in high-N addition plots than in low-N and control plots, regardless of its origin. In contrast, litter from high-N addition plots decomposing fastest in year 2, regardless of its destination. Finally, there was no effect of either litter origin or destination on the rate of decomposition in year 3. Litter collected in high-N addition plots had a concentration of N 12?17% higher than litter collected in other plots and higher in 2009 than in other years. Four years after the beginning of the fertilisation experiment, concentration and the microbial activity in the soil did not differ between the treatments. Our findings suggest that the levels of N addition predicted for Neotropical savannas can alter litter N concentrations and the process of litter decomposition, but that the direction and magnitude of these changes may be challenging to predict since that precipitation can influence the mechanisms regulating decomposition in the Cerrado.</span>
Stuart JEM, Holland-Moritz H, Lewis LR, Jean M, Miller SN, McDaniel SF, Fierer N, Ponciano JM, Mack MC (2020) Host Identity as a Driver of Moss-Associated N2 Fixation Rates in Alaska. Ecosystems.
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Read Publication<span>Moss-associated N2 fixation provides a substantial but heterogeneous input of new N to nutrient-limited ecosystems at high latitudes. In spite of the broad diversity of mosses found in boreal and Arctic ecosystems, the extent to which host moss identity drives variation in N2 fixation rates remains largely undetermined. We used 15N2 incubations to quantify the fixation rates associated with 34 moss species from 24 sites ranging from 60° to 68° N in Alaska, USA. Remarkably, all sampled moss genera fixed N2, including well-studied feather and peat mosses and genera such as Tomentypnum, Dicranum, and Polytrichum. The total moss-associated N2 fixation rates ranged from almost zero to 3.2 mg N m−2 d−1, with an average of 0.8 mg N m−2 d−1, based on abundance-weighted averages of all mosses summed for each site. Random forest models indicated that moss taxonomic family was a better predictor of rate variation across Alaska than any of the measured environmental factors, including site, pH, tree density, and mean annual precipitation and temperature. Consistent with this finding, mixed models showed that trends in N2 fixation rates among moss genera were consistent across biomes. We also found “hotspots” of high fixation rates in one-fourth of sampled sites. Our results demonstrated the importance of moss identity in influencing N2 fixation rates. This in turn indicates the potential utility of moss identity when making ecosystem N input predictions and exploring other sources of process rate variation.</span>
Tao X, Feng J, Yang Y, Wang G, Tian R, Fan F, Ning D, Bates CT, Hale L, Yuan MM, Wu L, Gao Q, Lei J, Schuur EAG, Yu J, Bracho R, Luo Y, Konstantinidis KT, Johnston ER, Cole JR, Penton CR, Tiedje JM, Zhou J (2020) Winter warming in Alaska accelerates lignin decomposition contributed by Proteobacteria. Microbiome 8(1): 84.
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Read Publication<span>In a warmer world, microbial decomposition of previously frozen organic carbon (C) is one of the most likely positive climate feedbacks of permafrost regions to the atmosphere. However, mechanistic understanding of microbial mediation on chemically recalcitrant C instability is limited; thus, it is crucial to identify and evaluate active decomposers of chemically recalcitrant C, which is essential for predicting C-cycle feedbacks and their relative strength of influence on climate change. Using stable isotope probing of the active layer of Arctic tundra soils after depleting soil labile C through a 975-day laboratory incubation, the identity of microbial decomposers of lignin and, their responses to warming were revealed.</span>
Terrer C, Jackson RB, Prentice IC, Keenan TF, Kaiser C, Vicca S, Fisher JB, Reich PB, Stocker BD, Hungate BA, Peñuelas J, McCallum I, Soudzilovskaia NA, Cernusak LA, Talhelm AF, Van Sundert K, Piao S, Newton PCD, Hovenden MJ, Blumenthal DM, Liu YY, Müller C, Winter K, Field CB, Viechtbauer W, Van Lissa CJ, Hoosbeek MR, Watanabe M, Koike T, Leshyk VO, Polley HW, Franklin O (2020) Author Correction: Nitrogen and phosphorus constrain the CO2 fertilization of global plant biomass. Nature Climate Change 10(7): 696-697.
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Read Publication<span>An amendment to this paper has been published and can be accessed via a link at the top of the paper.</span>
Turetsky MR, Abbott BW, Jones MC, Anthony KW, Olefeldt D, Schuur EAG, Grosse G, Kuhry P, Hugelius G, Koven C, Lawrence DM, Gibson C, Sannel ABK, McGuire AD (2020) Carbon release through abrupt permafrost thaw. Nature Geoscience 13(2): 138-143.
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Read Publication<span>The permafrost zone is expected to be a substantial carbon source to the atmosphere, yet large-scale models currently only simulate gradual changes in seasonally thawed soil. Abrupt thaw will probably occur in <20% of the permafrost zone but could affect half of permafrost carbon through collapsing ground, rapid erosion and landslides. Here, we synthesize the best available information and develop inventory models to simulate abrupt thaw impacts on permafrost carbon balance. Emissions across 2.5 million km2 of abrupt thaw could provide a similar climate feedback as gradual thaw emissions from the entire 18 million km2 permafrost region under the warming projection of Representative Concentration Pathway 8.5. While models forecast that gradual thaw may lead to net ecosystem carbon uptake under projections of Representative Concentration Pathway 4.5, abrupt thaw emissions are likely to offset this potential carbon sink. Active hillslope erosional features will occupy 3% of abrupt thaw terrain by 2300 but emit one-third of abrupt thaw carbon losses. Thaw lakes and wetlands are methane hot spots but their carbon release is partially offset by slowly regrowing vegetation. After considering abrupt thaw stabilization, lake drainage and soil carbon uptake by vegetation regrowth, we conclude that models considering only gradual permafrost thaw are substantially underestimating carbon emissions from thawing permafrost.</span>
Uhey DA, Hofstetter RW, Remke M, Vissa S, Haubensak KA (2020) Climate and vegetation structure shape ant communities along elevational gradients on the Colorado Plateau. Ecology and Evolution.
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Read Publication<span>Terrestrial animal communities are largely shaped by vegetation and climate. With climate also shaping vegetation, can we attribute animal patterns solely to climate? Our study observes ant community changes along climatic gradients (i.e., elevational gradients) within different habitat types (i.e., open and forest) on the Colorado Plateau in the southwestern United States. We sampled ants and vegetation along two elevational gradients spanning 1,132 m with average annual temperature and precipitation differences of 5.7°C and 645mm, respectively. We used regression analyses and structural equation modeling to compare the explanatory powers and effect sizes of climate and vegetation variables on ants. Climate variables had the strongest correlations and the largest effect sizes on ant communities, while vegetation composition, richness, and primary productivity had relatively small effects. Precipitation was the strongest predictor for most ant community metrics. Ant richness and abundance had a negative relationship with precipitation in forested habitats, and positive in open habitats. Our results show strong direct climate effects on ants with little or no effects of vegetation composition or primary productivity, but contrasting patterns between vegetation type (i.e., forested vs. open) with precipitation. This indicates vegetation structure can modulate climate responses of ant communities. Our study demonstrates climate-animal relationships may vary among vegetation types which can impact both findings from elevational studies and how communities will react to changes in climate.</span>
Vallejos R, Pérez J, Ellison AM, Richardson AD (2020) A spatial concordance correlation coefficient with an application to image analysis. Spatial Statistics 40:100405.
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Read Publication<span>In this work we define a spatial concordance coefficient for second-order stationary processes. This problem has been widely addressed in a non-spatial context, but here we consider a coefficient that for a fixed spatial lag allows one to compare two spatial sequences along a 45°line. The proposed coefficient was explored for the bivariate Matérn and Wendland covariance functions. The asymptotic normality of a sample version of the spatial concordance coefficient for an increasing domain sampling framework was established for the Wendland covariance function. To work with large digital images, we developed a local approach for estimating the concordance that uses local spatial models on non-overlapping windows. Monte Carlo simulations were used to gain additional insights into the asymptotic properties for finite sample sizes. As an illustrative example, we applied this methodology to two similar images of a deciduous forest canopy. The images were recorded with different cameras but similar fields-of-view and within minutes of each other. Our analysis showed that the local approach helped to explain a percentage of the non-spatial concordance and provided additional information about its decay as a function of the spatial lag.</span>
Walker AP, De Kauwe MG, (2020) Integrating the evidence for a terrestrial carbon sink caused by increasing atmospheric CO2. New Phytologist.
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Read Publication<span>Atmospheric carbon dioxide concentration ([CO2]) is increasing, which increases leaf-scale photosynthesis and intrinsic water-use efficiency. These direct responses have the potential to increase plant growth, vegetation biomass, and soil organic matter; transferring carbon from the atmosphere into terrestrial ecosystems (a carbon sink). A substantial global terrestrial carbon sink would slow the rate of [CO2] increase and thus climate change. However, ecosystem CO2-responses are complex or confounded by concurrent changes in multiple agents of global change and evidence for a [CO2]-driven terrestrial carbon sink can appear contradictory. Here we synthesise theory and broad, multi-disciplinary evidence for the effects of increasing [CO2] (iCO2) on the global terrestrial carbon sink. Evidence suggests a substantial increase in global photosynthesis since pre-industry. Established theory, supported by experiments, indicates that iCO2 is likely responsible for about half of the increase. Global carbon budgeting, atmospheric data, and forest inventories indicate a historical carbon sink, and these apparent iCO2-responses are high in comparison with experiments and theory. Plant mortality and soil carbon iCO2-responses are highly uncertain. In conclusion, a range of evidence supports a positive terrestrial carbon sink in response to iCO2, albeit with uncertain magnitude and strong suggestion of a role for additional agents of global change.</span>
Walker XJ, Baltzer JL, Bourgeau-Chavez L, Day NJ, Dieleman CM, Johnstone JF, Kane ES, Rogers BM, Turetsky MR, Veraverbeke S, Mack MC (2020) Patterns of Ecosystem Structure and Wildfire Carbon Combustion Across Six Ecoregions of the North American Boreal Forest. Frontiers in Forests and Global Change 3(87).
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Read Publication<span>Increases in fire frequency, extent, and severity are expected to strongly impact the structure and function of boreal forest ecosystems. An important function of the boreal forest is its ability to sequester and store carbon (C). Increasing disturbance from wildfires, emitting large amounts of C to the atmosphere, may create a positive feedback to climate warming. Variation in ecosystem structure and function throughout the boreal forest is important for predicting the effects of climate warming and changing fire regimes on C dynamics. In this study, we compiled data on soil characteristics, stand structure, pre-fire C pools, C loss from fire, and the potential drivers of these C metrics from 527 sites distributed across six ecoregions of North America’s western boreal forests. We assessed structural and functional differences between these fire-prone ecoregions using data from 417 recently burned sites (2004–2015) and estimated ecoregion-specific relationships between soil characteristics and depth from 167 of these sites plus an additional 110 sites (27 burned, 83 unburned). We found that northern boreal ecoregions were generally older, stored and emitted proportionally more belowground than aboveground C, and exhibited lower rates of C accumulation over time than southern ecoregions. We present ecoregion-specific estimates of depth-wise soil characteristics that are important for predicting C combustion from fire. As climate continues to warm and disturbance from wildfires increases, the C dynamics of these fire-prone ecoregions are likely to change with significant implications for the global C cycle and its feedbacks to climate change.</span>
Walker XJ, Rogers BM, Veraverbeke S, Johnstone JF, Baltzer JL, Barrett K, Bourgeau-Chavez L, Day NJ, de Groot WJ, Dieleman CM, Goetz S, Hoy E, Jenkins LK, Kane ES, Parisien MA, Potter S, Schuur EAG, Turetsky M, Whitman E, Mack MC (2020) Fuel availability not fire weather controls boreal wildfire severity and carbon emissions. Nature Climate Change 10(12): 1130 - 1136.
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Read Publication<span>Carbon (C) emissions from wildfires are a key terrestrial–atmosphere interaction that influences global atmospheric composition and climate. Positive feedbacks between climate warming and boreal wildfires are predicted based on top-down controls of fire weather and climate, but C emissions from boreal fires may also depend on bottom-up controls of fuel availability related to edaphic controls and overstory tree composition. Here we synthesized data from 417 field sites spanning six ecoregions in the northwestern North American boreal forest and assessed the network of interactions among potential bottom-up and top-down drivers of C emissions. Our results indicate that C emissions are more strongly driven by fuel availability than by fire weather, highlighting the importance of fine-scale drainage conditions, overstory tree species composition and fuel accumulation rates for predicting total C emissions. By implication, climate change-induced modification of fuels needs to be considered for accurately predicting future C emissions from boreal wildfires.</span>
Wu W, Dijkstra P, Dippold MA (2020) 13C analysis of fatty acid fragments by gas chromatography mass spectrometry for metabolic flux analysis. Geochimica et Cosmochimica Acta 284: 92-106.
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Read Publication<span>When multiple metabolic pathways lead to the same product, compound-specific isotope analysis may not provide enough information to quantify the activities of the contributing pathways. Instead, identification of where in the molecule the 13C is incorporated is required. Here we show how knowledge of position-specific 13C incorporation in fatty acids (FA) and FA fragments can be used to quantitatively estimate the fluxes through the central C metabolic network. We developed a method to measure 13C enrichment of FA and FA fragments (ethanoate, propionate) using electron impact GC–MS. We tested the accuracy and repeatability of the measurements using natural abundance and position-specific 13C labelled standards and FA extracted from Bacillus licheniformis and Pseudomonas fluorescens grown with labelled and unlabelled glucose. The molecular ions of FA generally reflected theoretical predictions of mass isotopomer distributions for natural abundance values, but that of the associated FA fragments deviated from expected values, likely associated with McLafferty rearrangements of hydrogen. After correction for naturally occurring isotopes, 13C enrichments of FA and FA fragments showed good agreement with expected isotope composition of FA standards (root mean square error < 0.044 at%; δ13C of ∼ 40‰), natural abundance and labelled glucose. The unsaturated FA extracted from P. fluorescens deviated from expected values likely associated with problems of co-elution and ion suppression and were excluded from analysis. The ratio of glucose-1-13C to glucose-3-13C incorporation into FA fragments was high for B. licheniformis, but low for P. fluorescens. Metabolic flux modelling based on the 13C enrichment of ethanoate and propionate fragments showed that B. licheniformis used Embden-Meyerhof-Parnas and pentose phosphate pathway (66% and 30%, respectively), whereas P. fluorescens utilized Entner-Doudoroff and pentose phosphate pathway (72% and 27%, respectively). FA fragment analysis is therefore a promising tool to study central C metabolic network activities of co-occurring groups of microbes in intact and complex environmental communities.</span>
Ye Y, Dai X, Fu X, Kou L, Luo Y, Jiang L, Wang H (2020) Differences in the rhizosphere effects among trees, shrubs and herbs in three subtropical plantations and their seasonal variations. European Journal of Soil Biology 100:103218.
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Read Publication<span>Unique soil properties in rhizosphere can affect plant growth and biogeochemical cycles of ecosystems. While rhizosphere has been widely investigated, little is known about differences in the rhizosphere effect (RE) between co-existing overstory trees and understory shrubs and herbs in forest ecosystems. In this study, we investigated REs on soil chemical properties of overstory trees and understory shrubs and herbs in forest plantations of southern China. Bulk soil and rhizospheres were sampled in April, July and December 2017. Soil pH, nitrate and ammonium nitrogen, dissolved organic carbon, available phosphorus, total carbon, total nitrogen, and total phosphorus were tested. The REs were defined as the ratios of the chemical properties of the rhizospheres to those of the bulk soil. Our results showed that pH was lower and nutrient contents were higher in the plant rhizospheres than the bulk soil. REs were generally larger in trees than understory plants. The REs were larger in July than April and December. Our findings indicated that the RE varied among plant life forms, species and sampling times, emphasizing the functional role of the RE of understory vegetation in subtropical forests.</span>
Zhai DL, Wang J, Thaler P, Luo Y, Xu J (2020) Contrasted effects of temperature during defoliation vs. refoliation periods on the infection of rubber powdery mildew (Oidium heveae) in Xishuangbanna, China. International Journal of Biometeorology 64(11): 1835-1845.
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Read Publication<span>Rubber powdery mildew caused by the foliar fungi Oidium heveae is one of the main diseases affecting rubber plantations (Hevea brasiliensis) worldwide. It is particularly serious in sub-optimal growing areas, such as Xishuangbanna in SW China. To prevent and control this disease, fungicides causing serious environmental problems are widely used. Strong correlations between the infection level and the temperature variables were reported previously, but they were related to monthly data that did not allow unraveling the patterns during the entire sensitive period. We correlated the infection level of powdery mildew of rubber trees recorded over 2003–2011 with antecedent 365 days daily temperature variables using partial least squares (PLS) regression. Our PLS regression results showed that the infection level of powdery mildew responded differently to the temperature variables of the defoliation and refoliation periods. Further analysis with Kriging interpolation showed that the infection level increased by 20% and 11%, respectively, per 1 °C rise of the daily maximum and mean temperature in the defoliation season, while it decreased by 8% and 10%, respectively, per 1 °C rise of the daily maximum and temperature difference in the refoliation season. This pattern was likely linked to the effects of temperature on leaf phenology. It seems highly possible that the infection level of powdery mildew increases, as increasing trends of maximum temperature and mean temperature during the defoliation continue.</span>
Zhang Q, Wei W, Chen L, Yang L, Chen HYH, Luo Y (2020) Soil Water Availability Drives Changes in Community Traits Along a Hydrothermal Gradient in Loess Plateau Grasslands. Rangeland Ecology & Management 73(2): 276-284.
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Read Publication<span>Plant functional traits can be used to predict ecosystem responses to climate gradients, yet precipitation explains very little variation for most traits. Soil water availability directly influences plant water uptake and thus may assist with the improvement of plant trait–water relationships. However, this promise remains poorly realized due to rare tests. Here, we provide the first study that attempts to link climate factors, vertical soil water availability, and community composition at a regional scale. Our study paired field-measured vertical soil available water (0–300 cm) and community functional composition at 46 herbaceous grassland sites along a steep hydrothermal gradient in the Loess Plateau of Central China. Community functional composition was expressed via community-weighted means of eight traits. Structural equation modeling was employed to evaluate the role of vertical soil available water content, controlled by precipitation and air temperature, in affecting plant community-weighted traits. We found that soil available water content at depths of 20–100 cm was typically responsible for mediating the effects of precipitation and air temperature on plant community composition. This emerged as the predominant factor to explain variations in grassland response traits, including leaf area, specific leaf area, and leaf dry matter content. These traits exhibited clear drought-induced shifts along soil desiccation gradients and responded to drier conditions by reducing leaf area/specific leaf area and increasing leaf dry matter content. Our findings rehighlighted soil water availability as the core driver that needs to be considered in the restoration and management of dryland ecosystems.</span>
Zhang Q, Wei W, Chen L, Yang L, Luo Y, Cai A (2020) Plant traits in influencing soil moisture in semiarid grasslands of the Loess Plateau, China. Science of The Total Environment 718: 137355.
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Read Publication<span>Large-scale vegetation restoration projects pose threats to water resource security in water-limited regions. Thus, the quantification of how vegetation cover affects soil moisture is of key importance to support effective restoration schemes in drylands. However, the current understanding of such effects remains poor. For this study, an in-situ vegetation-removal experiment was conducted at 36 herbaceous grassland sites having different community compositions and topographical conditions in two adjacent loess watersheds of the Loess Plateau, China. The effects of vegetation cover (vegetation effects) on soil moisture were analyzed across soil profiles (0–180 cm) and two growing seasons. Overall, 13 plant traits and 7 topographic and soil properties were employed to evaluate how community compositions modulated vegetation effects on soil moisture. The results showed that vegetation cover increased soil moisture in the surface layer (0–20 cm) by 6.81% during wet periods (semi-monthly rainfall >30 mm) relative to an in-situ unvegetated control, but primarily induced a decline of soil moisture in the deep soil layer (20–180 cm) by 19.44% across two growing seasons. Redundancy analysis (RDA) and structural equation modeling (SEM) suggested that these vegetation effects on soil moisture were significantly correlated with vegetative height, leaf area, shallow root allocation, and slope gradient. Our study revealed that tall, small-leaved, and shallow-rooted plants on flat topographies were beneficial to soil water retention and replenishment. This implied that current restoration strategies may be significantly improved through the development of optimal communities and diverse terracing measures. Our findings are anticipated to provide effective guidance for soil water conservation, as well as ecosystem rehabilitation in dry and degraded regions.</span>
Zheng M, Chen H, Li D, Luo Y, Mo J (2020) Substrate stoichiometry determines nitrogen fixation throughout succession in southern Chinese forests. Ecology Letters 23(2): 336-347.
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Read Publication<span>The traditional view holds that biological nitrogen (N) fixation often peaks in early- or mid-successional ecosystems and declines throughout succession based on the hypothesis that soil N richness and/or phosphorus (P) depletion become disadvantageous to N fixers. This view, however, fails to support the observation that N fixers can remain active in many old-growth forests despite the presence of N-rich and/or P-limiting soils. Here, we found unexpected increases in N fixation rates in the soil, forest floor, and moss throughout three successional forests and along six age-gradient forests in southern China. We further found that the variation in N fixation was controlled by substrate carbon(C) : N and C : (N : P) stoichiometry rather than by substrate N or P. Our findings highlight the utility of ecological stoichiometry in illuminating the mechanisms that couple forest succession and N cycling.</span>
Zheng M, Zhou Z, Zhao P, Luo Y, Ye Q, Zhang K, Song L, Mo J (2020) Effects of human disturbance activities and environmental change factors on terrestrial nitrogen fixation. Global Change Biology 26(11): 6203-6217.
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Read Publication<span>Biological nitrogen (N) fixation plays an important role in terrestrial N cycling and represents a key driver of terrestrial net primary productivity (NPP). Despite the importance of N fixation in terrestrial ecosystems, our knowledge regarding the controls on terrestrial N fixation remains poor. Here, we conducted a meta-analysis (based on 852 observations from 158 studies) of N fixation across three types of ecosystems with different status of disturbance (no management, restoration [previously disturbed], and disturbance [currently disturbed]) and in response to multiple environmental change factors (warming, elevated carbon dioxide [CO2], increased precipitation, increased drought, increased N deposition, and their combinations). We explored the mechanisms underlying the changes in N fixation by examining the variations in soil physicochemical properties (bulk density, texture, moisture, and pH), plant and microbial characteristics (dominant plant species numbers, plant coverage, and soil microbial biomass), and soil resources (total carbon, total N, total phosphorus (P), inorganic N, and inorganic P). Human disturbance inhibited non-symbiotic N fixation but not symbiotic N fixation. Terrestrial N fixation was stimulated by warming (+152.7%), elevated CO2 (+19.6%), and increased precipitation (+73.1%) but inhibited by increased drought (?30.4%), N deposition (?31.0%), and combinations of available multiple environmental change factors (?14.5%), the extents of which varied among biomes and ecosystem compartments. Human disturbance reduced the N fixation responses to environmental change factors, which was associated with the changes in soil physicochemical properties (2%?56%, p < .001) and the declines in plant and microbial characteristics (3%?49%, p ≤ .003) and soil resources (6%?48%, p ≤ .03). Overall, our findings reveal for the first time the effects of multiple environmental change factors on terrestrial N fixation and indicate the role of human disturbance activities in inhibiting N fixation, which can improve our understanding, modeling, and prediction of terrestrial N budgets, NPP, and ecosystem feedbacks under global change scenarios.</span>
Zhou Z, Wang C, and Luo Y (2020) Meta-analysis of the impacts of global change factors on soil microbial diversity and functionality. Nature Communications 11(1):3072.
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Read Publication<span>Biodiversity on the Earth is changing at an unprecedented rate due to a variety of global change factors (GCFs). However, the effects of GCFs on microbial diversity is unclear despite that soil microorganisms play a critical role in biogeochemical cycling. Here, we synthesize 1235 GCF observations worldwide and show that microbial rare species are more sensitive to GCFs than common species, while GCFs do not always lead to a reduction in microbial diversity. GCFs-induced shifts in microbial alpha diversity can be predominately explained by the changed soil pH. In addition, GCF impacts on soil functionality are explained by microbial community structure and biomass rather than the alpha diversity. Altogether, our findings of GCF impacts on microbial diversity are fundamentally different from previous knowledge for well-studied plant and animal communities, and are crucial to policy-making for the conservation of microbial diversity hotspots under global changes.</span>
2019
Bian H, Geng Q, Xiao H, Shen C, Li Q, Cheng X, Luo Y, Ruan H, Xu X (2019) Fine Root Biomass Mediates Soil Fauna Community in Response to Nitrogen Addition in Poplar Plantations (Populus deltoids) on the East Coast of China. Forests 10(2): 122.
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Read PublicationSoil fauna is critical for maintaining ecosystem functioning, and its community could be significantly impacted by nitrogen (N) deposition. However, our knowledge of how soil-faunal community composition responds to N addition is still limited. In this study, we simulated N deposition (0, 50, 100, 150, and 300 kg N ha-1 year-1) to explore the effects of N addition on the total and the phytophagous soil fauna along the soil profile (0-10, 10-25, and 25-40 cm) in poplar plantations (Populus deltoids) on the east coast of China. Ammonium nitrate (NH4NO3) was dissolved in water and sprayed evenly under the canopy with a backpack sprayer to simulate N deposition. Our results showed that N addition either significantly increased or decreased the density (D) of both the total and the phytophagous soil fauna (Dtotal and Dp) at low or high N addition rates, respectively, indicating the existence of threshold effects over the range of N addition. However, N addition had no significant impacts on the number of groups (G) and diversity (H) of either the total or the phytophagous soil fauna (Gtotal, Gp and Htotal, Hp). With increasing soil depth, Dtotal, Dp, Gtotal, and Gp largely decreased, showing that the soil fauna have a propensity to aggregate at the soil surface. Htotal and Hp did not significantly vary along the soil profile. Importantly, the threshold effects of N addition on Dtotal and Dp increased from 50 and 100 to 150 kg N ha-1 year-1 along the soil profile. Fine root biomass was the dominant factor mediating variations in Dtotal and Dp. Our results suggested that N addition may drive changes in soil-faunal community composition by altering belowground food resources in poplar plantations.
Boyd MA, Berner LT, Doak P, Goetz S, Rogers B, Wagner D, Walker X, Mack MC (2019) Impacts of climate and insect herbivory on productivity and physiology of trembling aspen (Populus tremuloides) in Alaskan boreal forests. Environmental Research Letters.
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Read PublicationClimate change is impacting forested ecosystems worldwide, particularly in the Northern Hemisphere where warming has increased at a faster rate than the rest of the globe. As climate warms, trembling aspen (Populus tremuloides) is expected to become more successful in northern boreal forests because of its current presence in drier areas of North America. However, large-scale productivity decline of aspen has recently been documented throughout the United States and Canada as a result of drought and insect outbreaks. We used tree ring measurements (basal area increment (BAI) and stable carbon isotopes (δ13C)) and remote sensing indices of vegetation productivity (NDVI) to study the impact of climate and damage by the aspen epidermal leaf miner (Phyllocnistis populiella) on aspen productivity and physiology in interior Alaska. We found that productivity decreased with greater leaf mining and was not sensitive to growing season moisture availability. Although productivity decreased during high leaf mining years, it recovered to pre-outbreak levels during years of low insect damage, suggesting a degree of resilience to P. populiella mining. Climate and leaf mining interacted to influence tree ring δ13C, with greater leaf mining resulting in decreased δ13C when growing season moisture availability was low. We also found that NDVI was negatively associated with leaf mining, and positively correlated with BAI and the δ13C decrease corresponding to mining. This suggests that NDVI is capturing not only variations in productivity, but also changes in physiology associated with P. populiella. Overall, these findings indicate that the indirect effects of P. populiella mining have a larger impact on aspen productivity and physiology than climate under current conditions, and is essential to consider when assessing growth, physiology and NDVI trends in interior Alaska.
Butterfield BJ, Anderson RS, Holmgren CA, Betancourt JL (2019) Extinction debt and delayed colonization have had comparable but unique effects on plant community–climate lags since the Last Glacial Maximum. Global Ecology and Biogeography 28(8): 1067-1077.
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Read PublicationPlant communities typically exhibit lagged responses to climate change due to poorly understood effects of colonization and local extinction. Here, we quantify rates of change in mean cold tolerances, and contributions of colonization and local extinction to those rates, recorded in plant macrofossil assemblages from North American hot deserts over the last 30,000 years. Location: Mojave, Sonoran and Chihuahuan Deserts. Time period 30-0 thousand years before present (kybp). Major taxa studied: Vascular plants. Methods: Colonization and local extinction dates for 269 plant species were approximated from macrofossils in 15 packrat (Neotoma) midden series. Cold tolerances estimated from contemporary climate were used to quantify assemblage-mean cold tolerances through time. Rates of colonization and local extinction, and their effects on rates of change in assemblage-mean cold tolerances, were estimated for 30-20 kybp (Late Pleistocene, no directional warming), 20-10 kybp (deglaciation, rapid warming) and 10-0 kybp (Holocene, no directional warming). Results: Rates of change in all metrics were negligible during the Late Pleistocene. Rates of change in assemblage-mean cold tolerances (mean 1.0°C x 10-4/yr) lagged behind warming during deglaciation, and continued at similar rates (1.2°C x 10-4/yr) throughout the Holocene. Colonization and local extinction contributed equally to delayed responses to warming, but their dynamics differed through time: Colonization by warm-adapted species predominated during deglaciation, while the most heat-adapted species exhibited long delays in colonization. Only the most cold-adapted species went locally extinct during deglaciation, followed by slow repayment of the extinction debt of cool-adapted species during the Holocene. Main conclusions Responses to rapid warming can persist for millennia, even after cessation of warming. Consistent patterns from different midden series across the region support a metacommunity model in which dispersal interacts with environmental filters and buffers against local extinction to drive community?climate disequilibrium during and after periods of warming.
Butterfield BJ, Holmgren CA, Anderson RS, Betancourt JL (2019) Life history traits predict colonization and extinction lags of desert plant species since the Last Glacial Maximum. Ecology e02817.
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Read PublicationVariation in life-history strategies can affect metapopulation dynamics and consequently the composition and diversity of communities. However, data sets that allow for the full range of species turnover from colonization to extinction over relevant time periods are limited. The late Quaternary record provides unique opportunities to explore the traits that may have influenced interspecific variation in responses to past climate warming, in particular the rate at which species colonized newly suitable habitat or went locally extinct from degrading habitat. We controlled for differences in species climate niches in order to predict expected colonization and extinction sequences recorded in packrat middens from 15 localities in the Mohave, Sonoran, and Chihuahuan deserts of North America. After accounting for temperature niche differences, we tested the hypotheses that dispersal syndrome (none, wind, vertebrate), growth form (herb, shrub, tree) and seed mass mediated variation in postglacial colonization lags among species, whereas clonality (clonal, non-clonal), growth form, and seed mass affected extinction lags. Growth form and dispersal syndrome interactively affected colonization lags, where herbaceous species lacking long-distance dispersal mechanisms exhibited lags that exceeded those of woody, wind or vertebrate-dispersed species by an average of 2,000-5,000 yr. Growth form and seed mass interactively affected extinction lags, with very small-seeded shrubs persisting for 4,000-8,000 yr longer than other functional groups. Taller, vertebrate-dispersed plants have been shown in other studies to disperse farther than shorter plants without specialized dispersal mechanisms. We found that variation along this axis of dispersal syndromes resulted in dramatic differences in colonization rates in response to past climate change. Very small seeded shrubs may have a unique combination of long vegetative and seed bank lifetimes that may allow them to persist for long periods despite declines in habitat condition. This study indicates that readily measurable traits may help predict which species will be more or less sensitive to future climate change, and inform interventions that can stabilize and promote at-risk populations.
Cai A, Chang N, Zhang W, Liang G, Zhang X, Hou E, Jiang L, Chen X, Xu M, Luo Y (2019) The spatial patterns of litter turnover time in Chinese terrestrial ecosystems. European Journal of Soil Science .
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<div class="coolBar__section">The feedback between plant, soil and climate is partly determined by plant litter turnover time, which is influenced by climate, litter quality and soil properties. However, the spatial patterns of litter turnover time and its interrelation with these variables are rarely quantified. With a database of 1,378 litter turnover times and key associated climate, litter quality and soil properties (total of 20 variables), this study investigated the driving factors and spatial patterns of litter turnover time across Chinese terrestrial ecosystems. The mean litter turnover time was the longest in forest ecosystems, followed by that in grassland and cropland ecosystems. The litter turnover time varied significantly depending on the litter quality and climate zone, and increased exponentially as latitude increased. Mean annual temperature (MAT) and mean annual precipitation (MAP) could accurately predict litter turnover time via negative exponential equations. Among these variables, MAT had the greatest influence on litter turnover time, which accounted for 37.4% of the variation, followed by litter quality (ecosystem types, litter types, C:N of litter and lignin content; 33.4%) and soil properties (sand content, soil pH and soil organic carbon (SOC); 29.2%) based on a boosted regression tree (BRT) model. Path analysis identified that MAT negatively affected litter turnover time both directly and indirectly through regulating soil properties and litter quality, which positively and directly affected litter turnover time. Finally, the spatial patterns of litter turnover time were obtained with a regional dataset of ecosystem types, MAT, sand content, soil pH and SOC as BRT model drivers. Overall, our results suggest that climate variables have contrasting effects on litter turnover time and could mediate the impact on litter turnover time by litter quality and soil properties. These results highlight important implications for climate‐smart soil management and can be used to create reliable model predictions.</div>
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Cai A, Xu M, Wang B, Zhang W, Liang G, Hou E, Luo Y (2019) Manure acts as a better fertilizer for increasing crop yields than synthetic fertilizer does by improving soil fertility. Soil and Tillage Research 189: 168-175.
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Read PublicationFertilization is an important management strategy for crop yields by mediating soil fertility. However, rare studies quantitatively assessed the interactions among fertilization, crop yields, and soil fertility. Here, data from a 25-year fertilization experiment in the humid subtropical region of Southern China were used to evaluate and quantify the effect of fertilization on crop yields via soil fertility. Seven treatments were chosen: CK (non-fertilizer); N (synthetic nitrogen); NP (synthetic N and phosphorus); NPK (synthetic N, P and potassium); NPKM1 (synthetic NPK with manure); 1.5NPKM1 (1.5 times of NPKM1); and M2 (manure alone). Overall, the crop yields of wheat and maize under manure (1.36–1.58 and 3.85-5.82 Mg ha−1) were higher than those under CK (0.34 and 0.25 Mg ha−1) and synthetic fertilized treatments (0.27–0.97 and 0.48–2.65 Mg ha−1), as the averaged of 1991–2015. Higher SOC stocks were found under the NPKM1, 1.5NPKM1, and M2 treatments with a pronounced increase in SOC over the first 10 years and stable over the last 15 years. By the boosted regression trees, manure, synthetic fertilizer and soil properties (SOC storage, soil pH, and soil nutrients) accounted for 39%, 21%, and 40% of the variation of the relative yield, respectively. Path analysis identified a network of inter-relations of manure, synthetic fertilizer, and soil properties in the relative yields. Compared to synthetic fertilized treatments, manure application strongly and positively affected the relative yield by increasing SOC storage, soil nutrients, and soil pH (path coefficients: 0.90, 0.88, and 0.76). These factors explained 72% of the crop yields' variance. These results suggest that manure application is a viable strategy for regulating crop yields due to its improvement in soil fertility.
Carbone MS, Seyednasrollah B, Rademacher TT, Basler D, Le Moine JM, Beals S, Beasley J, Greene A, Kelroy J, Richardson AD (2019) Flux Puppy – An open-source software application and portable system design for low-cost manual measurements of CO2 and H2O fluxes. Agricultural and Forest Meteorology 274:1-6.
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Read PublicationManual chamber-based measurements of CO2 (and H2O) fluxes are important for understanding ecosystem carbon metabolism. Small opaque chambers can be used to measure leaf, stem and soil respiration. Larger transparent chambers can be used to measure net ecosystem exchange of CO2, and small jars often serve this purpose for laboratory incubations of soil and plant material. We developed an Android application (app), called Flux Puppy, to facilitate chamber-based flux measurements in the field and laboratory. The app is designed to run on an inexpensive handheld Android device, such as a tablet or phone, and it has a graphical user interface that communicates with a LI-COR LI-820 and LI-830 (CO2) or LI-840 and LI-850 (CO2/H2O) infrared gas analyzer. The app logs concentrations of CO2 and H2O, cell temperature and pressure at 1 Hz, displays the output graphically, and calculates the linear regression slope, R-squared, and standard error of the CO2 time series. A metadata screen allows users to enter operator, site, and plot information, as well as take a photograph using the Android device’s built-in camera, and log measurement location using the device GPS. Additionally, there is a notes field, which can be revised after the measurements are taken. Data files (the 1 s raw data, photograph, and metadata including statistics calculated from the raw data) are then transmitted off the device through file sharing options (Gmail, Outlook, Google Drive, Dropbox etc.). Because Flux Puppy code is open-source (available on GitHub) and the flux measurement system we describe is relatively inexpensive and straightforward to assemble, it should be of broad interest to the carbon cycling community.
Chen J, Zhang Y, Luo Y, Zhou X, Jiang Y, Zhao J, Chen Y, Wang C, Guo L, Cao J (2019) Contrasting responses after fires of the source components of soil respiration and ecosystem respiration. European Journal of Soil Science 70(3): 616-629.
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Read PublicationWildfire is an important ecological disturbance that can have cascading effects on ecosystem carbon (C) fluxes. Ecosystem respiration (ER) and soil respiration (SR) account for two of the largest terrestrial C fluxes to the atmosphere, and they play critical roles in regulating C?climate feedbacks. Here, the responses of ER, SR and their source components to experimental burning in a meadow grassland on the Tibetan Plateau were investigated. Fire treatment increased ER by 9% but decreased SR by 15%. The contrasting post-fire responses of SR and ER can be explained by the behaviour of their source components; that is, fire increased aboveground plant respiration (Ragb) by 37%, but decreased heterotrophic respiration (HR) by 21%. Increases in ER and Ragb were mainly related to enhanced plant productivity, whereas smaller SR and HR were associated with reductions in microbial biomass and soil moisture. Accounting for the responses of ER, SR and their intrinsic components has advanced our understanding of how fire affects ecosystem C fluxes. Highlights Fire treatment increased ecosystem respiration (ER) and aboveground plant respiration. Fire treatment decreased soil respiration (SR) and heterotrophic respiration (HR). Increases in ER and aboveground plant respiration were related to plant productivity. Reductions in SR and HR were caused by the suppressed microbial activity.
Chen Y, Chen J, Luo Y (2019) Data-driven ENZYme (DENZY) model represents soil organic carbon dynamics in forests impacted by nitrogen deposition. Soil Biology and Biochemistry 138: 107575.
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Read PublicationSoil microorganisms participate in almost all soil organic carbon (SOC) transformations, but they are not represented explicitly in the current generation of earth system models. This study used a data-driven approach to incorporate extracellular enzyme activity into the Terrestrial ECOsystem (TECO) model, and the updated version was named the Data-driven ENZYme (DENZY) model. DENZY is based on results from an extensive data synthesis, which show that the CN ratio is positively correlated with ligninase activity (R2 = 0.50). The latter is inversely correlated to soil organic carbon storage. The DENZY model was parameterized using the revise database to information from a recent meta-analysis and tested for its ability to simulate SOC dynamics at Duke Forest (North Carolina, USA) from 1996 to 2007. DENZY can well simulate the observed negative relationship between ligninase activity and SOC under N deposition conditions (R2 ranges from 0.61 to 0.89). Moreover, outputs from DENZY better matched the observed SOC than its prototype model with the same parameterization. This study provides a simple and straightforward approach to effectively use real-world observations to improve SOC projections in terrestrial biogeochemical models.
Copeland SM, Munson SM, Bradford JB, Butterfield BJ, Gunnell KL (2019) Long-term plant community trajectories suggest divergent responses of native and non-native perennials and annuals to vegetation removal and seeding treatments. Restoration Ecology.
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Read PublicationLand managers frequently apply vegetation removal and seeding treatments to restore ecosystem function following woody plant encroachment, invasive species spread, and wildfire. However, the long-term outcome of these treatments is unclear due to a lack of widespread monitoring. We quantified how vegetation removal (via wildfire or management) with or without seeding and environmental conditions related to plant community composition change over time in 491 sites across the intermountain western United States. Most community metrics took over 10 years to reach baseline conditions posttreatment, with the slowest recovery observed for native perennial cover. Total cover was initially higher in sites with seeding after vegetation removal than sites with vegetation removal alone, but increased faster in sites with vegetation removal only. Seeding after vegetation removal was associated with rapidly increasing non-native perennial cover and decreasing non-native annual cover. Native perennial cover increased in vegetation removal sites irrespective of seeding and was suppressed by increasing non-native perennial cover. Seeding was associated with higher non-native richness across the monitoring period as well as initially higher, then declining, total and native species richness. Several cover and richness recovery metrics were positively associated with mean annual precipitation and negatively associated with mean annual temperature, whereas relationships with weather extremes depended on the lag time and season. Our results suggest that key plant groups, such as native perennials and non-native annuals, respond to restoration treatments at divergent timescales and with different sensitivities to climate and weather variation.
Day NJ, Dunfield KE, Johnstone JF, Mack MC, Turetsky MR, Walker XJ, White AL, Baltzer JL (2019) Wildfire severity reduces richness and alters composition of soil fungal communities in boreal forests of western Canada. Global Change Biology.
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Read PublicationWildfire is the dominant disturbance in boreal forests and fire activity is increasing in these regions. Soil fungal communities are important for plant growth and nutrient cycling postfire but there is little understanding of how fires impact fungal communities across landscapes, fire severity gradients, and stand types in boreal forests. Understanding relationships between fungal community composition, particularly mycorrhizas, and understory plant composition is therefore important in predicting how future fire regimes may affect vegetation. We used an extreme wildfire event in boreal forests of Canada's Northwest Territories to test drivers of fungal communities and assess relationships with plant communities. We sampled soils from 39 plots 1 year after fire and 8 unburned plots. High-throughput sequencing (MiSeq, ITS) revealed 2,034 fungal operational taxonomic units. We found soil pH and fire severity (proportion soil organic layer combusted), and interactions between these drivers were important for fungal community structure (composition, richness, diversity, functional groups). Where fire severity was low, samples with low pH had higher total fungal, mycorrhizal, and saprotroph richness compared to where severity was high. Increased fire severity caused declines in richness of total fungi, mycorrhizas, and saprotrophs, and declines in diversity of total fungi and mycorrhizas. The importance of stand age (a surrogate for fire return interval) for fungal composition suggests we could detect long-term successional patterns even after fire. Mycorrhizal and plant community composition, richness, and diversity were weakly but significantly correlated. These weak relationships and the distribution of fungi across plots suggest that the underlying driver of fungal community structure is pH, which is modified by fire severity. This study shows the importance of edaphic factors in determining fungal community structure at large scales, but suggests these patterns are mediated by interactions between fire and forest stand composition.
Fang XM, Zhang XL, Chen FS, Zong YY, Bu WS, Wan SZ, Luo Y, Wang H (2019) Phosphorus addition alters the response of soil organic carbon decomposition to nitrogen deposition in a subtropical forest. Soil Biology and Biochemistry 133: 119-128.
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Read PublicationThe continuous increase of nitrogen (N) deposition may exacerbate phosphorus (P) deficiency, which affects soil organic carbon (SOC) decomposition by changing microbial community characteristics in subtropical forests with highly weathered soils. However, there is currently little information about the role of P and the N × P interaction in SOC dynamics. Here, a field nutrient manipulation experiment was established in a subtropical plantation forest in China. Soils collected from simulated N deposition and P addition treatments for 5 years were incubated at 25 °C for 130 days. Soil microbial composition was measured using the phospholipid fatty acid method and the enzyme activities related to SOC hydrolysis were measured. The SOC concentration and δ13C in bulk soil and three particle-size fracfractions were also determined. The cumulative CO2 respired over 9 days, representing the utilization of carbon sources under field conditions, increased with N deposition levels under the without-P treatment, while no significant differences were found among the three N deposition levels in the with-P treatment. Meanwhile, P addition generally suppressed the SOC decomposition during 130 days incubation. Similarly, P addition decreased the potential organic carbon decomposition (C0) and C0/SOC ratio. In contrast, C0 increased with N deposition in the without-P treatment, while was unaffected by N deposition under the with-P treatment, suggesting the response of SOC decomposition to N deposition was affected following P addition by alteration of SOC quality. Moreover, N deposition tended to deplete the δ13C of the SOC and P addition enriched the δ13C of the macro-particulate organic carbon. Addition of P increased total microbial, fungal and bacterial biomass values by 41.6%, 90.0% and 46.9%, respectively, whereas N deposition had no significant effect. Soil fungi/bacteria ratio significantly increased by N deposition and P addition, which partly explained the reduction of SOC decomposition after P addition. The cellobioside activity significantly decreased by 48.3% after P addition, while cellobioside and β-xylosidase activities increased with N deposition, suggesting that N deposition and P addition had opposite roles in the SOC stability. These results indicate that the positive effect of N deposition on SOC decomposition was suppressed when P was added by changing microbial community and enzyme activity and enhanced P availability may result in increased SOC accumulation under N deposition scenarios in subtropical forests.
Feng J, Penton CR, He Z, Van Nostrand JD, Yuan MM, Wu L, Wang C, Qin Y, Shi Z, Guo X, Schuur EAG, Luo Y, Bracho R, Konstantinidis KT, Cole JR, Tiedje JM, Yang Y, Zhou J (2019) Long-term warming in Alaska enlarges the diazotrophic community in deep soils. mBio 10(1): e02521-18.
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Read PublicationTundra ecosystems are typically carbon (C) rich but nitrogen (N) limited. Since biological N2 fixation is the major source of biologically available N, the soil N2-fixing (i.e., diazotrophic) community serves as an essential N supplier to the tundra ecosystem. Recent climate warming has induced deeper permafrost thaw and adversely affected C sequestration, which is modulated by N availability. Therefore, it is crucial to examine the responses of diazotrophic communities to warming across the depths of tundra soils. Herein, we carried out one of the deepest sequencing efforts of nitrogenase gene (nifH) to investigate how 5 years of experimental winter warming affects Alaskan soil diazotrophic community composition and abundance spanning both the organic and mineral layers. Although soil depth had a stronger influence on diazotrophic community composition than warming, warming significantly (P &amp;lt; 0.05) enhanced diazotrophic abundance by 86.3% and aboveground plant biomass by 25.2%. Diazotrophic composition in the middle and lower organic layers, detected by nifH sequencing and a microarray-based tool (GeoChip), was markedly altered, with an increase of α-diversity. Changes in diazotrophic abundance and composition significantly correlated with soil moisture, soil thaw duration, and plant biomass, as shown by structural equation modeling analyses. Therefore, more abundant diazotrophic communities induced by warming may potentially serve as an important mechanism for supplementing biologically available N in this tundra ecosystem.IMPORTANCE With the likelihood that changes in global climate will adversely affect the soil C reservoir in the northern circumpolar permafrost zone, an understanding of the potential role of diazotrophic communities in enhancing biological N2 fixation, which constrains both plant production and microbial decomposition in tundra soils, is important in elucidating the responses of soil microbial communities to global climate change. A recent study showed that the composition of the diazotrophic community in a tundra soil exhibited no change under a short-term (1.5-year) winter warming experiment. However, it remains crucial to examine whether the lack of diazotrophic community responses to warming is persistent over a longer time period as a possibly important mechanism in stabilizing tundra soil C. Through a detailed characterization of the effects of winter warming on diazotrophic communities, we showed that a long-term (5-year) winter warming substantially enhanced diazotrophic abundance and altered community composition, though soil depth had a stronger influence on diazotrophic community composition than warming. These changes were best explained by changes in soil moisture, soil thaw duration, and plant biomass. These results provide crucial insights into the potential factors that may impact future C and N availability in tundra regions.
Foster AC, Armstrong AH, Shuman JK, Shugart HH, Rogers BM, Mack MC, Goetz SJ, Ranson KJ (2019) Importance of tree- and species-level interactions with wildfire, climate, and soils in interior Alaska: Implications for forest change under a warming climate. Ecological Modeling 409: 108765.
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Read PublicationThe boreal zone of Alaska is dominated by interactions between disturbances, vegetation, and soils. These interactions are likely to change in the future through increasing permafrost thaw, more frequent and intense wildfires, and vegetation change from drought and competition. We utilize an individual tree-based vegetation model, the University of Virginia Forest Model Enhanced (UVAFME), to estimate current and future forest conditions across sites within interior Alaska. We updated UVAFME for application within interior Alaska, including improved simulation of permafrost dynamics, litter decay, nutrient dynamics, fire mortality, and post-fire regrowth. Following these updates, UVAFME output on species-specific biomass and stem density was comparable to inventory measurements at various forest types within interior Alaska. We then simulated forest response to climate change at specific inventory locations and across the Tanana Valley River Basin on a 2 × 2 km2 grid. We derived projected temperature and precipitation from a five-model average taken from the CMIP5 archive under the RCP 4.5 and 8.5 scenarios. Results suggest that climate change and the concomitant impacts on wildfire and permafrost dynamics will result in overall decreases in biomass (particularly for spruce (Picea spp.)) within the interior Tanana Valley, despite increases in quaking aspen (Populus tremuloides) biomass, and a resulting shift towards higher deciduous fraction. Simulation results also predict increases in biomass at cold, wet locations and at high elevations, and decreases in biomass in dry locations, under both moderate (RCP 4.5) and extreme (RCP 8.5) climate change scenarios. These simulations demonstrate that a highly detailed, species interactive model can be used across a large region within Alaska to investigate interactions between vegetation, climate, wildfire, and permafrost. The vegetation changes predicted here have the capacity to feed back to broader scale climate-forest interactions in the North American boreal forest, a region which contributes significantly to the global carbon and energy budgets.
Ge R, He H, Ren X, Zhang L, Yu G, Smallman TL, Zhou T, Yu SY, Luo Y, Xie Z, Wang S, Wang H, Zhou G, Zhang Q, Wang A, Fan Z, Zhang Y, Shen W, Yin H, Lin L (2019) Underestimated ecosystem carbon turnover time and sequestration under the steady state assumption: A perspective from long-term data assimilation. Global Change Biology 25(3): 938-953.
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Read PublicationIt is critical to accurately estimate carbon (C) turnover time as it dominates the uncertainty in ecosystem C sinks and their response to future climate change. In the absence of direct observations of ecosystem C losses, C turnover times are commonly estimated under the steady state assumption (SSA), which has been applied across a large range of temporal and spatial scales including many at which the validity of the assumption is likely to be violated. However, the errors associated with improperly applying SSA to estimate C turnover time and its covariance with climate as well as ecosystem C sequestrations have yet to be fully quantified. Here, we developed a novel model-data fusion framework and systematically analyzed the SSA-induced biases using time-series data collected from 10 permanent forest plots in the eastern China monsoon region. The results showed that (a) the SSA significantly underestimated mean turnover times (MTTs) by 29%, thereby leading to a 4.83-fold underestimation of the net ecosystem productivity (NEP) in these forest ecosystems, a major C sink globally; (b) the SSA-induced bias in MTT and NEP correlates negatively with forest age, which provides a significant caveat for applying the SSA to young-aged ecosystems; and (c) the sensitivity of MTT to temperature and precipitation was 22% and 42% lower, respectively, under the SSA. Thus, under the expected climate change, spatiotemporal changes in MTT are likely to be underestimated, thereby resulting in large errors in the variability of predicted global NEP. With the development of observation technology and the accumulation of spatiotemporal data, we suggest estimating MTTs at the disequilibrium state via long-term data assimilation, thereby effectively reducing the uncertainty in ecosystem C sequestration estimations and providing a better understanding of regional or global C cycle dynamics and C-climate feedback.
Geyer KM, Dijkstra P, Sinsabaugh R, Frey SD (2019) Clarifying the interpretation of carbon use efficiency in soil through methods comparison. Soil Biology and Biochemistry 128: 79-88.
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Read PublicationAccurate estimates of microbial carbon use efficiency (CUE) are required to predict how global change will impact microbially-mediated ecosystem functions such as organic matter decomposition. Multiple approaches are currently used to quantify CUE but the extent to which estimates reflect methodological variability is unknown. This limits our ability to apply or cross-compare published CUE values. Here we evaluated the performance of five methods in a single soil under standard conditions. The microbial response to three substrate amendment rates (0.0, 0.05, and 2.0 mg glucose-C g−1 soil) was examined using: 13C and 18O isotope tracing approaches which estimate CUE based on substrate uptake and growth dynamics; calorespirometry which infers growth and CUE from metabolic heat and respiration rates; metabolic flux analysis where CUE is determined as the balance between biosynthesis and respiration using position-specific 13CO2 production of labeled glucose; and stoichiometric modeling which derives CUE from elemental ratios of microbial biomass, substrate, and exoenzyme activity. The CUE estimates we obtained differed by method and substrate concentration, ranging under in situ conditions from <0.4 for the substrate-nonspecific methods that do not use C tracers (18O, stoichiometric modeling) to >0.6 for the substrate-specific methods that trace glucose use (13C method, calorespirometry, metabolic flux analysis). We explore the different aspects of microbial metabolism that each method captures and how this affects the interpretation of CUE estimates. We recommend that users consider the strengths and weaknesses of each method when choosing the technique that will best address their research needs.
Grove S, Saarman NP, Gilbert GS, Faircloth B, Haubensak KA, Parker IM (2019) Ectomycorrhizas and tree seedling establishment are strongly influenced by forest edge proximity but not soil inoculum. Ecological Applications e01867.
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Read PublicationReforestation is challenging when timber harvested areas have been degraded, invaded by nonnative species, or are of marginal suitability to begin with. Conifers form mutualistic partnerships with ectomycorrhizal fungi (EMF) to obtain greater access to soil resources, and these partnerships may be especially important in degraded areas. However, timber harvest can impact mycorrhizal fungi by removing or compacting topsoil, removing host plants, and warming and drying the soil. We used a field experiment to evaluate the role of EMF in Douglas-fir reforestation in clearcuts invaded by Cytisus scoparius (Scotch broom) where traditional reforestation approaches have repeatedly failed. We tested how planting distance from intact Douglas-fir forest edges influenced reforestation success and whether inoculation with forest soils can be used to restore EMF relationships. We used an Illumina DNA sequencing approach to measure the abundance, richness and composition of ectomycorrhizal fungi on Douglas-fir roots, and assessed differences in Douglas-fir seedling survival and growth near to and far from forest edges with and without forest soil inoculum. Planting Douglas-fir seedlings near forest edges increased seedling survival, growth, and EMF root colonization. Edge proximity had no effect on EMF richness but did change fungal community composition. Inoculations with forest soil did not increase EMF abundance or richness or change community composition, nor did it improve seedling establishment. With Illumina sequencing, we identified two to three times greater species richness than described in previous edge effects studies. Of the 95 EMF species we identified, 40% of the species occurred on less than 5% of the seedlings. The ability to detect fungi at low abundance may explain why we did not detect differences in EMF richness with distance to hosts as previous studies. Our findings suggest that forest edges are suitable for reforestation, even when the interiors of deforested areas are not. We advocate for timber harvest designs that maximize edge habitat where ectomycorrhizal fungi contribute to tree establishment. However, this study does not support the use of inoculation with forest soil as a simple method to enhance EMF and seedling survival.
Guerrieri R, Belmecheri S, Ollinger SV, Asbjornsen H, Jennings K, Xiao J, Stocker BD, Martin M, Hollinger DY, Bracho-Garrillo R, Clark K, Dore S, Kolb T, Munger JW, Novick K, Richardson AD (2019) Disentangling the role of photosynthesis and stomatal conductance on rising forest water-use efficiency. Proceedings of the National Academy of Sciences 116(34): 16909.
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Read PublicationForests remove about 30% of anthropogenic CO2 emissions through photosynthesis and return almost 40% of incident precipitation back to the atmosphere via transpiration. The trade-off between photosynthesis and transpiration through stomata, the water-use efficiency (WUE), is an important driver of plant evolution and ecosystem functioning, and has profound effects on climate. Using stable carbon and oxygen isotope ratios in tree rings, we found that WUE has increased by a magnitude consistent with estimates from atmospheric measurements and model predictions. Enhanced photosynthesis was widespread, while reductions in stomatal conductance were modest and restricted to moisture-limited forests. This result points to smaller reductions in transpiration in response to increasing atmospheric CO2, with important implications for forest–climate interactions, which remain to be explored.Multiple lines of evidence suggest that plant water-use efficiency (WUE)—the ratio of carbon assimilation to water loss—has increased in recent decades. Although rising atmospheric CO2 has been proposed as the principal cause, the underlying physiological mechanisms are still being debated, and implications for the global water cycle remain uncertain. Here, we addressed this gap using 30-y tree ring records of carbon and oxygen isotope measurements and basal area increment from 12 species in 8 North American mature temperate forests. Our goal was to separate the contributions of enhanced photosynthesis and reduced stomatal conductance to WUE trends and to assess consistency between multiple commonly used methods for estimating WUE. Our results show that tree ring-derived estimates of increases in WUE are consistent with estimates from atmospheric measurements and predictions based on an optimal balancing of carbon gains and water costs, but are lower than those based on ecosystem-scale flux observations. Although both physiological mechanisms contributed to rising WUE, enhanced photosynthesis was widespread, while reductions in stomatal conductance were modest and restricted to species that experienced moisture limitations. This finding challenges the hypothesis that rising WUE in forests is primarily the result of widespread, CO2-induced reductions in stomatal conductance.
Guo JS, Ogle K (2019) Antecedent soil water content and vapor pressure deficit interactively control water potential in Larrea tridentata. New Phytologist 221(1): 218-232.
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Read PublicationPlant water potential Ψ is regulated by stomatal responses to atmospheric moisture demand D and soil water availability W, but the timescales of influence and interactions between these drivers of plant Ψ are poorly understood. Here, we quantify the effects of antecedent D and W on plant Ψ in the desert shrub Larrea tridentata. Repeated measurements of plant baseline water potential ΨB and diurnal water potential ΨD were analyzed in a Bayesian framework to evaluate the influence of antecedent D and W at daily and subdaily timescales. Both ΨB and ΨD exhibited negative, 2- to 4-d lagged responses to daily-scale D; conversely, plant ΨD responded almost instantaneously to subdaily D, though the direction of this response depended on antecedent moisture conditions. Plant ΨB and ΨD responded positively and immediately (no lag) to shallow W, which contrasts the negative, lagged (6-7 d) response to deep W. The changing sensitivity of ΨD to subdaily D highlights shifting modes of plant Ψ regulation: D effects on ΨD range from negative to neutral to positive depending on past conditions and time of day. Explicit consideration of antecedent conditions across multiple timescales can reveal important complexities in plant responses.
Guo X, Zhou X, Hale L, Yuan M, Ning D, Feng J, Shi Z, Li Z, Feng B, Gao Q, Wu L, Shi W, Zhou A, Fu Y, Wu L, He Z, Van Nostrand JD, Qiu G, Liu X, Luo Y, Tiedje JM, Yang Y, Zhou J (2019) Climate warming accelerates temporal scaling of grassland soil microbial biodiversity. Nature Ecology & Evolution 3(4): 612-619.
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Read PublicationDetermining the temporal scaling of biodiversity, typically described as species–time relationships (STRs), in the face of global climate change is a central issue in ecology because it is fundamental to biodiversity preservation and ecosystem management. However, whether and how climate change affects microbial STRs remains unclear, mainly due to the scarcity of long-term experimental data. Here, we examine the STRs and phylogenetic–time relationships (PTRs) of soil bacteria and fungi in a long-term multifactorial global change experiment with warming (+3 °C), half precipitation (−50%), double precipitation (+100%) and clipping (annual plant biomass removal). Soil bacteria and fungi all exhibited strong STRs and PTRs across the 12 experimental conditions. Strikingly, warming accelerated the bacterial and fungal STR and PTR exponents (that is, the w values), yielding significantly (P < 0.001) higher temporal scaling rates. While the STRs and PTRs were significantly shifted by altered precipitation, clipping and their combinations, warming played the predominant role. In addition, comparison with the previous literature revealed that soil bacteria and fungi had considerably higher overall temporal scaling rates (w = 0.39–0.64) than those of plants and animals (w = 0.21–0.38). Our results on warming-enhanced temporal scaling of microbial biodiversity suggest that the strategies of soil biodiversity preservation and ecosystem management may need to be adjusted in a warmer world.
Hale L, Feng W, Yin H, Guo X, Zhou X, Brach R, Pegoraro E, Penton CR, Wu L, Cole J, Konstantinidis KT, Luo Y, Tiedje JM, Schuur EAG, Zhou J (2019) Tundra microbial community taxa and traits predict decomposition parameters of stable, old soil organic carbon. The ISME Journal 13(12): 2901-2915.
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Read PublicationThe susceptibility of soil organic carbon (SOC) in tundra to microbial decomposition under warmer climate scenarios potentially threatens a massive positive feedback to climate change, but the underlying mechanisms of stable SOC decomposition remain elusive. Herein, Alaskan tundra soils from three depths (a fibric O horizon with litter and course roots, an O horizon with decomposing litter and roots, and a mineral-organic mix, laying just above the permafrost) were incubated. Resulting respiration data were assimilated into a 3-pool model to derive decomposition kinetic parameters for fast, slow, and passive SOC pools. Bacterial, archaeal, and fungal taxa and microbial functional genes were profiled throughout the 3-year incubation. Correlation analyses and a Random Forest approach revealed associations between model parameters and microbial community profiles, taxa, and traits. There were more associations between the microbial community data and the SOC decomposition parameters of slow and passive SOC pools than those of the fast SOC pool. Also, microbial community profiles were better predictors of model parameters in deeper soils, which had higher mineral contents and relatively greater quantities of old SOC than in surface soils. Overall, our analyses revealed the functional potential of microbial communities to decompose tundra SOC through a suite of specialized genes and taxa. These results portray divergent strategies by which microbial communities access SOC pools across varying depths, lending mechanistic insights into the vulnerability of what is considered stable SOC in tundra regions.
Hilman B, Muhr J, Trumbore SE, Kunert N, Carbon MS, Yuval P, Wright SJ, Moreno G, Perez-Priego O, Migliavacca M, Carrara A, Grunzweig YO, Weiner T, Angert A (2019) Comparison of CO2 and O2 fluxes demonstrate retention of respired CO2 in tree stems from a range of tree species. Biogeosciences 16: 177-191.
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Read PublicationThe ratio of CO efflux to influx (ARQ, apparent respiratory quotient) in tree stems is expected to be 1.0 for carbohydrates, the main substrate supporting stem respiration. In previous studies of stem fluxes, ARQ values below 1.0 were observed and hypothesized to indicate retention of respired carbon within the stem. Here, we demonstrate that stem ARQ 1.0 values are common across 85 tropical, temperate, and Mediterranean forest trees from nine different species. Mean ARQ values per species per site ranged from 0.39 to 0.78, with an overall mean of 0.59. Assuming that uptake provides a measure of in situ stem respiration (due to the low solubility of O2), the overall mean indicates that on average 41% of CO2 respired in stems is not emitted from the local stem surface. The instantaneous ARQ did not vary with sap flow. ARQ values of incubated stem cores were similar to those measured in stem chambers on intact trees. We therefore conclude that dissolution of CO2 in the xylem sap and transport away from the site of respiration cannot explain the low ARQ values. We suggest refixation of respired CO2 in biosynthesis reactions as possible mechanism for low ARQ values.
Hou E, Lu X, Jiang L, Wen D, Luo Y (2019) Quantifying soil phosphorus dynamics: A data assimilation approach. Journal of Geophysical Research: Biogeosciences 124(7): 2159-2173.
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Read PublicationThe dynamics of soil phosphorus (P) control its bioavailability. Yet it remains a challenge to quantify soil P dynamics. Here we developed a soil P dynamics (SPD) model. We then assimilated eight data sets of 426-day changes in Hedley P fractions into the SPD model, to quantify the dynamics of six major P pools in eight soil samples that are representative of a wide type of soils. The performance of our SPD model was better for labile P, secondary mineral P, and occluded P than for nonoccluded organic P (Po) and primary mineral P. All parameters describing soil P dynamics were approximately constrained by the data sets. The average turnover rates were labile P 0.040 g g?1 day?1, nonoccluded Po 0.051 g g?1 day?1, secondary mineral P 0.023 g g?1 day?1, primary mineral P 0.00088 g g?1 day?1, occluded Po 0.0066 g g?1 day?1, and occluded inorganic P 0.0065 g g?1 day?1, in the greenhouse environment studied. Labile P was transferred on average more to nonoccluded Po (transfer coefficient of 0.42) and secondary mineral P (0.38) than to plants (0.20). Soil pH and organic C concentration were the key soil properties regulating the competition for P between plants and soil secondary minerals. The turnover rate of labile P was positively correlated with that of nonoccluded Po and secondary mineral P. The pool size of labile P was most sensitive to its turnover rate. Overall, we suggest data assimilation can contribute significantly to an improved understanding of soil P dynamics.
Huang Y, Stacy M, Jiang J, Sundi N, Ma S, Saruta V, Jung CG, Shi Z, Xia J, Hanson PJ, Ricciuto D, Luo Y (2019) Realized ecological forecast through an interactive Ecological Platform for Assimilating Data (EcoPAD, v1.0) into models. Geoscientific Model Development 12: 1119-1137.
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Read Publication<p id="d1e248">Predicting future changes in ecosystem services is not only highly desirable but is also becoming feasible as several forces (e.g., available big data, developed data assimilation (DA) techniques, and advanced cyber-infrastructure) are converging to transform ecological research into quantitative forecasting. To realize ecological forecasting, we have developed an Ecological Platform for Assimilating Data (EcoPAD, v1.0) into models. EcoPAD (v1.0) is a web-based software system that automates data transfer and processing from sensor networks to ecological forecasting through data management, model simulation, data assimilation, forecasting, and visualization. It facilitates interactive data–model integration from which the model is recursively improved through updated data while data are systematically refined under the guidance of model. EcoPAD (v1.0) relies on data from observations, process-oriented models, DA techniques, and the web-based workflow.</p>
<p id="d1e251">We applied EcoPAD (v1.0) to the Spruce and Peatland Responses Under Climatic and Environmental change (SPRUCE) experiment in northern Minnesota. The EcoPAD-SPRUCE realizes fully automated data transfer, feeds meteorological data to drive model simulations, assimilates both manually measured and automated sensor data into the Terrestrial ECOsystem (TECO) model, and recursively forecasts the responses of various biophysical and biogeochemical processes to five temperature and two <span class="inline-formula">CO<sub>2</sub></span> treatments in near-real time (weekly). Forecasting with EcoPAD-SPRUCE has revealed that mismatches in forecasting carbon pool dynamics are more related to model (e.g., model structure, parameter, and initial value) than forcing variables, opposite to forecasting flux variables. EcoPAD-SPRUCE quantified acclimations of methane production in response to warming treatments through shifted posterior distributions of the <span class="inline-formula">CH<sub>4</sub>:CO<sub>2</sub></span> ratio and the temperature sensitivity (<span class="inline-formula"><i>Q</i><sub>10</sub></span>) of methane production towards lower values. Different case studies indicated that realistic forecasting of carbon dynamics relies on appropriate model structure, correct parameterization, and accurate external forcing. Moreover, EcoPAD-SPRUCE stimulated active feedbacks between experimenters and modelers to identify model components to be improved<span id="page1120"></span> and additional measurements to be taken. It has become an interactive model–experiment (ModEx) system and opens a novel avenue for interactive dialogue between modelers and experimenters. Altogether, EcoPAD (v1.0) acts to integrate multiple sources of information and knowledge to best inform ecological forecasting.</p>
Hurteau Matthew D, North Malcolm P, Koch George W, Hungate Bruce A (2019) Opinion: Managing for disturbance stabilizes forest carbon. Proceedings of the National Academy of Sciences 116(21).
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Read PublicationForest ecosystems sequester approximately 12% of anthropogenic carbon emissions, and efforts to increase forest carbon uptake are central to climate change mitigation policy. Managing forests to store carbon has focused on increasing forested area, decreasing area lost to logging and clearing, and increasing forest carbon density. Warming, drought, and wildfires challenge the stability of carbon stored in forests. By contrast, natural cycles of low-intensity fires in dry forests can, over the long term, promote forest carbon storage by protecting carbon in soil and in large, old trees. The conundrum is how to balance immediate, disturbance-driven carbon loss with long-term, stable carbon storage and account for these risks in policies for forest carbon management.
Hutchings JA, Bianchi TS, Kaufman DS, Kholodov AL, Vaughn DR, Schuur EAG (2019) Millennial-scale carbon accumulation and molecular transformation in a permafrost core from Interior Alaska. Geochimica et Cosmochimica Acta 253: 231-248.
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Read PublicationOrganic carbon stored in high-latitude permafrost represents a potential positive feedback to climate warming as well as a valuable store of paleoenvironmental information. The below-freezing conditions have effectively removed permafrost organic material from the modern carbon cycle and preserved its pre-freezing bulk and molecular states. The conditions that lead to efficient burial of organic carbon (OC) within permafrost were investigated by measuring OC stocks, past accumulation rates, and biogeochemical composition of a permafrost core taken from Interior Alaska dating back to 40 ka. The post-glacial Marine Isotope Stage 1 is represented by the top 1.2 m of the core and contains 64.7 kg OC/m2 with an accumulation rate of 4.3 g OC/m2/yr. The sediments that accumulated around the Last Glacial Maximum contain 9.9 kg OC/m2 with an accumulation rate of 0.5 g OC/m2/yr. Carbon storage (144.7 kg OC/m2) and accumulation (26.1 g OC/m2/yr) are both observed to be greatest between 35 and 40 ka, late during the Marine Isotope Stage 3 global interstadial. The extent of OC degradation was assessed using lignin and amino acid biomarkers with both approaches indicating well-preserved contemporary active layer and interstadial OC, whereas stadial OC was highly degraded. Lignin compositional indices throughout the core appear altered by sorptive processes that confounded some expected trends in the overall organic matter composition, while amino acids provided a more integrated pattern of change. Significant correlations between carbon-normalized hydroxyproline and total lignin concentrations further support the usefulness of hydroxyproline as an indicator for the abundance of plant organic matter. A novel amino acid plant-microbial index of the ratio of microbial-specific muramic acid and diaminopimelic acid biomarkers to the plant-specific hydroxyproline biomarker, indicate a transition from plant-dominated organic matter in fresh organic soils (index values of 0.01–0.20) to more microbial-dominated organic matter in degraded mineral soils (index values of 0.50–2.50). The branched glycerol dialkyl glycerol tetraether composition is complex and is not immediately compatible with existing temperature transfer functions. Residence time within the active layer is interpreted to integrate key factors such as primary productivity, inorganic sediment delivery, and other climate factors that control soil organic matter degradation. The Marine Isotope Stage 3, mid-Wisconsin interstadial period at this locality was forest-dominated and suggests the currently prevailing tundra ecotone is sensitive to environmental change. The majority of buried permafrost OC is high in degradability and if thawed, would be expected to be highly vulnerable to microbial decomposition.
Jackson CR, Stone BWG (2019) Canopy position is a stronger determinant of bacterial community composition and diversity than environmental disturbance in the phyllosphere. FEMS Microbology Ecology 95(4): fiz032.
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Read PublicationThe effect of rain on the phyllosphere community has not been extensively explored, especially in the context of spatial variation on the impact of rain throughout the tree canopy. We characterized the response of the phyllosphere bacterial community removed from leaf surfaces of the Southern Magnolia (Magnolia grandiflora) to rain across different spatial locations of the canopy. We hypothesized that: (1) rain would lead to an initial decrease in phyllosphere bacterial diversity, followed by an increase in diversity on subsequent days, but that this effect would be minimized in the lower and interior portion of the canopy, and that (2) community beta dispersion of phyllosphere microorganisms would be lower following rain, and similarly contingent on canopy position. We used targeted next-generation sequencing of the V4 region of the bacterial 16S rRNA gene to characterize bacterial composition. We found higher bacterial richness in interior canopy and distinct composition across canopy positions. Further, the effect of rain on beta dispersion was contingent on canopy position: rain lowered dispersion in the upper canopy but increased it in the lower and interior canopy. Our results demonstrate that canopy structure should be considered when looking at the impact of rain on the collective phyllosphere community.
Jiang D, Geng Q, Li Q, Luo Y, Vogel J, Shi Z, Ruan H, Xu X (2019) Nitrogen and phosphorus resorption in planted forests worldwide. Forests 10(3): 201.
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Read PublicationNutrient resorption from senescing leaves is one of the plants&rsquo; essential nutrient conservation strategies. Parameters associated with resorption are important nutrient-cycling constraints for accurate predictions of long-term primary productivity in forest ecosystems. However, we know little about the spatial patterns and drivers of leaf nutrient resorption in planted forests worldwide. By synthesizing results of 146 studies, we explored nitrogen (N) and phosphorus (P) resorption efficiency (NRE and PRE) among climate zones and tree functional types, as well as the factors that play dominant roles in nutrient resorption in plantations globally. Our results showed that the mean NRE and PRE were 58.98% &plusmn; 0.53% and 60.21% &plusmn; 0.77%, respectively. NRE significantly increased from tropical to boreal zones, while PRE did not significantly differ among climate zones, suggesting differential impacts of climates on NRE and PRE. Plant functional types exert a strong influence on nutrient resorption. Conifer trees had higher PRE than broadleaf trees, reflecting the adaptation of the coniferous trees to oligotrophic habitats. Deciduous trees had lower PRE than evergreen trees that are commonly planted in P-limited low latitudes and have long leaf longevity with high nutrient use efficiency. While non-N-fixing trees had higher NRE than N-fixing trees, the PRE of non-N-fixing trees was lower than that of N-fixing trees, indicating significant impact of the N-fixing ability on the resorption of N and P. Our multivariate regression analyses showed that variations in NRE were mainly regulated by climates (mean annual precipitation and latitude), while variations in PRE were dominantly controlled by green leaf nutrient concentrations (N and P). Our results, in general, suggest that the predicted global warming and changed precipitation regimes may profoundly affect N cycling in planted forests. In addition, green leaf nutrient concentrations may be good indicators for PRE in planted forests.
Jiang Y, Qian H, Wang L, Feng J, Huang S, Hungate BA, van Kessel C, Horwath WR, Zhang X, Quin X, Li Y, Feng X, Zhang J, Deng A, Zheng C, Song Z, Hu S, van Groenigen KJ, Zhang W (2019) Limited potential of harvest index improvement to reduce methane emissions from rice paddies. Global Change Biology 25(2): 686-698.
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Read PublicationRice is a staple food for nearly half of the world's population, but rice paddies constitute a major source of anthropogenic CH4 emissions. Root exudates from growing rice plants are an important substrate for methane-producing microorganisms. Therefore, breeding efforts optimizing rice plant photosynthate allocation to grains, i.e., increasing harvest index (HI), are widely expected to reduce CH4 emissions with higher yield. Here we show, by combining a series of experiments, meta-analyses and an expert survey, that the potential of CH4 mitigation from rice paddies through HI improvement is in fact small. Whereas HI improvement reduced CH4 emissions under continuously flooded (CF) irrigation, it did not affect CH4 emissions in systems with intermittent irrigation (II). We estimate that future plant breeding efforts aimed at HI improvement to the theoretical maximum value will reduce CH4 emissions in CF systems by 4.4%. However, CF systems currently make up only a small fraction of the total rice growing area (i.e., 27% of the Chinese rice paddy area). Thus, to achieve substantial CH4 mitigation from rice agriculture, alternative plant breeding strategies may be needed, along with alternative management.
Johnston ER, Hatt JK, Wu L, Guo X, Luo Y, Schuur EAG, Tiedje JM, Zhou J, Konstantinidis KT (2019) Responses of tundra soil microbial communities to half a decade of experimental warming at two critical depths. Proceedings of the National Academy of Sciences 116(30): 15096.
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Read PublicationOngoing permafrost thaw is expected to stimulate microbial release of greenhouse gases, threatening to further exacerbate climate change (cause positive feedback). In this study, a unique field warming experiment was conducted in Interior Alaska to promote surface permafrost degradation while maintaining uniform hydraulic conditions. After 5 winters of experimental warming by ∼1 °C, microbial community shifts were observed at the receded permafrost/active layer boundary, which reflected more reduced conditions, including increased methanogenesis. In contrast, increased carbohydrate utilization (respiration) was observed at the surface layer. These shifts were relatable to observed increases in CO2 and CH4 release from this study site and the surrounding ecosystem. Collectively, our results demonstrate that microbial responses to warming are rapid and identify potential biomarkers that could be important in modeling.Northern-latitude tundra soils harbor substantial carbon (C) stocks that are highly susceptible to microbial degradation with rising global temperatures. Understanding the magnitude and direction (e.g., C release or sequestration) of the microbial responses to warming is necessary to accurately model climate change. In this study, Alaskan tundra soils were subjected to experimental in situ warming by ∼1.1 °C above ambient temperature, and the microbial communities were evaluated using metagenomics after 4.5 years, at 2 depths: 15 to 25 cm (active layer at outset of the experiment) and 45 to 55 cm (transition zone at the permafrost/active layer boundary at the outset of the experiment). In contrast to small or insignificant shifts after 1.5 years of warming, 4.5 years of warming resulted in significant changes to the abundances of functional traits and the corresponding taxa relative to control plots (no warming), and microbial shifts differed qualitatively between the two soil depths. At 15 to 25 cm, increased abundances of carbohydrate utilization genes were observed that correlated with (increased) measured ecosystem carbon respiration. At the 45- to 55-cm layer, increased methanogenesis potential was observed, which corresponded with a 3-fold increase in abundance of a single archaeal clade of the Methanosarcinales order, increased annual thaw duration (45.3 vs. 79.3 days), and increased CH4 emissions. Collectively, these data demonstrate that the microbial responses to warming in tundra soil are rapid and markedly different between the 2 critical soil layers evaluated, and identify potential biomarkers for the corresponding microbial processes that could be important in modeling.
Jung CG, Xu X, Niu S, Liang J, Chen X, Shi Z, Jiang L, Luo Y (2019) Experimental warming amplified opposite impacts of drought vs. wet extremes on ecosystem carbon cycle in a tallgrass prairie. Agricultural and Forest Meteorology 276-277: 107635.
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Read PublicationClimate warming is leading to greater precipitation variability, resulting in increased frequency and intensity of both drought and wet extremes. However, how these extreme events interact with climate warming and hay-harvest in grasslands to impact ecosystem functions has not yet been well explored. In this study, we took advantage of a long-term experiment to examine how climate warming and clipping (i.e., mimicking hay harvest) regulated impacts of naturally occurring drought and wet extremes on ecosystem CO2 fluxes of a tallgrass prairie in the Great Plains, USA. Warming resulted in net ecosystem carbon release (i.e., positive net ecosystem CO2 exchange, NEE) in the extreme drought year of 2011, but significantly enhanced net carbon uptake in the extremely wet year of 2015 in comparison with NEE in normal years. Warming-induced carbon release in the drought year was due to significantly enhanced ecosystem respiration (ER) from mid-summer to early-autumn, whereas warming-enhanced NEE in the wet year was due to an increase in aboveground net primary production (ANPP) compared to those in normal years. Drought diminished warming-induced increases in ANPP to about one sixth of that in the wet year in the unclipped plots. Interestingly, clipping offset the drought-mediated ecosystem carbon loss by increasing GPP and weakened the wet-enhanced ANPP. Overall, our results suggest that a future, warmer climate may exacerbate carbon losses in terrestrial ecosystems during drought extremes but stimulate the ecosystem carbon sink under wet extremes.
Knox SH, Jackson RB, Poulter B, McNicol G, Fluet-Chouinard E, Zhang Z, Hugelius G, Bousquet P, Canadell JG, Saunois M, Papale D, Chu H, Keenan TF, Baldocchi D, Torn MS, Mammarella I, Trotta C, Aurela M, Bohrer G, Campbell DI, Cescatti A, Chamberlain S, Chen J, Chen W, Dengel S, Desai AR, Euskirchen E, Friborg T, Gasbarra D, Goded I, Goeckede M, Heimann M, Helbig M, Hirano T, Hollinger DY, Iwata H, Kang M, Klatt J, Krauss KW, Kutzbach L, Lohila A, Mitra B, Morin TH, Nilsson MB, Niu S, Noormets A, Oechel WC, Peichl M, Peltola O, Reba ML, Richardson AD, Runkle BRK, RY Y, Sachs T, Schafer KVR, Schmid HP, Shurpali N, Sonnentag O, Tang ACI, Ueyama M, Vargas R, Vesala T, Ward EJ, Windham-Myers L, Wohlfahrt G, Zona D (2019) FLUXNET-CH4 synthesis activity: Objectives, observations, and future directions. Bulletin of the American Meteorological Society.
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Read PublicationKomatsu KJ, Avolio ML, Lemoine NP, Isbell F, Grman E, Houseman GR, Koerner SE, Johnson DS, Wilcox KR, Alatalo JM, Anderson JP, Aerts R, Baer SG, Baldwin AH, Bates J, Beierkuhnlein C, Belote RT, Blair J, Bloor JMG, Bohlen PJ, Bork EW, Boughton EH, Bowman WD, Britton AJ, Cahill JF, Chaneton E, Chiariello NR, Cheng J, Collins, SL, Cornelissen JHC, Du G, Eskelinen A, Firn J, Foster B, Gough L, Gross K, Hallett LM, Han X, Harmens H, Hovenden MJ, Jagerbrand A, Jentsch A, Kern C, Klanderud K, Knapp AK, Kreyling J, Li W, Luo Y, McCulley, RL, McLaren JR, Megonigal JP, Morgan JW, Onipchenko V, Pennings, SC, Prevéy JS, Price JN, Reich, PB, Robinson CH, Russell FL, Sala OE, Seabloom EW, Smith MD, Soudzilovskaia NA, Souza L, Suding K, Suttle KB, Svejcar T, Tilman D, Tognetti P, Turkington R, White S, Xu Z, Yahdjian L, Yu Q, Zhang P, Zhang Y (2019) Global change effects on plant communities are magnified by time and the number of global change factors imposed. Proceedings of the National Academy of Sciences 116(36): 17867.
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Read PublicationAccurate prediction of community responses to global change drivers (GCDs) is critical given the effects of biodiversity on ecosystem services. There is consensus that human activities are driving species extinctions at the global scale, but debate remains over whether GCDs are systematically altering local communities worldwide. Across 105 experiments that included over 400 experimental manipulations, we found evidence for a lagged response of herbaceous plant communities to GCDs caused by shifts in the identities and relative abundances of species, often without a corresponding difference in species richness. These results provide evidence that community responses are pervasive across a wide variety of GCDs on long-term temporal scales and that these responses increase in strength when multiple GCDs are simultaneously imposed.Global change drivers (GCDs) are expected to alter community structure and consequently, the services that ecosystems provide. Yet, few experimental investigations have examined effects of GCDs on plant community structure across multiple ecosystem types, and those that do exist present conflicting patterns. In an unprecedented global synthesis of over 100 experiments that manipulated factors linked to GCDs, we show that herbaceous plant community responses depend on experimental manipulation length and number of factors manipulated. We found that plant communities are fairly resistant to experimentally manipulated GCDs in the short term (&amp;lt;10 y). In contrast, long-term (≥10 y) experiments show increasing community divergence of treatments from control conditions. Surprisingly, these community responses occurred with similar frequency across the GCD types manipulated in our database. However, community responses were more common when 3 or more GCDs were simultaneously manipulated, suggesting the emergence of additive or synergistic effects of multiple drivers, particularly over long time periods. In half of the cases, GCD manipulations caused a difference in community composition without a corresponding species richness difference, indicating that species reordering or replacement is an important mechanism of community responses to GCDs and should be given greater consideration when examining consequences of GCDs for the biodiversity–ecosystem function relationship. Human activities are currently driving unparalleled global changes worldwide. Our analyses provide the most comprehensive evidence to date that these human activities may have widespread impacts on plant community composition globally, which will increase in frequency over time and be greater in areas where communities face multiple GCDs simultaneously.
Kwon MJ, Natali SM, Hicks Pries CE, Schuur EAG, Steinhof A, Crummer KG, Zimov N, Zimov SA, Heimann M, Kolle O, Gockede M (2019) Drainage enhances modern soil carbon contribution but reduces old soil carbon contribution to ecosystem respiration in tundra ecosystems. Global Change Biology.
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Read PublicationWarming temperatures are likely to accelerate permafrost thaw in the Arctic, potentially leading to the release of old carbon previously stored in deep frozen soil layers. Deeper thaw depths in combination with geomorphological changes due to the loss of ice structures in permafrost, may modify soil water distribution, creating wetter or drier soil conditions. Previous studies revealed higher ecosystem respiration rates under drier conditions, and this study investigated the cause of the increased ecosystem respiration rates using radiocarbon signatures of respired CO2 from two drying manipulation experiments: one in moist and the other in wet tundra. We demonstrate that higher contributions of CO2 from shallow soil layers (0?15 cm; modern soil carbon) drive the increased ecosystem respiration rates, while contributions from deeper soil (below 15 cm from surface and down to the permafrost table; old soil carbon) decreased. These changes can be attributed to more aerobic conditions in shallow soil layers, but also the soil temperature increases in shallow layers but decreases in deep layers, due to the altered thermal properties of organic soils. Decreased abundance of aerenchymatous plant species following drainage in wet tundra reduced old carbon release but increased aboveground plant biomass elevated contributions of autotrophic respiration to ecosystem respiration. The results of this study suggest that drier soils following drainage may accelerate decomposition of modern soil carbon in shallow layers but slow down decomposition of old soil carbon in deep layers, which may offset some of the old soil carbon loss from thawing permafrost. This article is protected by copyright. All rights reserved.
Lantz TC, Moffat ND, Fraser RH, Walker X (2019) Reproductive limitation mediates the response of white spruce (Picea glauca) to climate warming across the forest–tundra ecotone. Arctic Science 5(4): 167-184.
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Read PublicationShifts in the extent of the boreal forest during past warm intervals and correlations between climate and the position of the forest–tundra ecotone suggest that recent temperature increases will facilitate forest expansion into tundra ecosystems. In this study, we used a unique set of high-resolution repeat photographs to characterize white spruce (<i>Picea glauca</i> (Moench) Voss) populations in 1980 and 2015 at 52 sites across the forest–tundra transition in the Northwest Territories, Canada. We also conducted field inventories at eight sites to examine mapping accuracy, construct age distributions, and assess cone production and seed viability. Our analysis shows that stand density in the forest–tundra has increased significantly since 1980 but that the density of spruce at sites in the tundra has not changed. Age distributions constructed from field sampling also indicate that recent recruitment has occurred in the forest–tundra but not at tundra sites. The nonlinear relationship between summer temperature and seed viability suggests that recent warming has facilitated recruitment in the northern Subarctic but that cold temperatures still limit recruitment at higher latitude tundra sites. Additional research to determine the extent of changes in forest density across the northern Subarctic should be conducted to determine if similar changes are occurring across this ecotone.
Li J, Mau RL, Dijkstra P, Koch BJ, Schwartz E, Liu XJA, Morrissey EM, Blazewicz SJ, Pett-Ridge J, Stone BW, Hayer M, Hungate BA (2019) Predictive genomic traits for bacterial growth in culture versus actual growth in soil. The ISME Journal.
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Read PublicationRelationships between microbial genes and performance are often evaluated in the laboratory in pure cultures, with little validation in nature. Here, we show that genomic traits related to laboratory measurements of maximum growth potential failed to predict the growth rates of bacteria in unamended soil, but successfully predicted growth responses to resource pulses: growth increased with 16S rRNA gene copy number and declined with genome size after substrate addition to soils, responses that were repeated in four different ecosystems. Genome size best predicted growth rate in response to addition of glucose alone; adding ammonium with glucose weakened the relationship, and the relationship was absent in nutrient-replete pure cultures, consistent with the idea that reduced genome size is a mechanism of nutrient conservation. Our findings demonstrate that genomic traits of soil bacteria can map to their ecological performance in nature, but the mapping is poor under native soil conditions, where genomic traits related to stress tolerance may prove more predictive. These results remind that phenotype depends on environmental context, underscoring the importance of verifying proposed schemes of trait-based strategies through direct measurement of performance in nature, an important and currently missing foundation for translating microbial processes from genes to ecosystems.
Li J, Wang G, Mayes MA, Allison SD, Frey SD, Shi Z, Hu XM, Luo Y, Melillo JM (2019) Reduced carbon use efficiency and increased microbial turnover with soil warming. Global Change Biology 25(3): 900-910.
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Read PublicationGlobal soil carbon (C) stocks are expected to decline with warming, and changes in microbial processes are key to this projection. However, warming responses of critical microbial parameters such as carbon use efficiency (CUE) and biomass turnover (rB) are not well understood. Here, we determine these parameters using a probabilistic inversion approach that integrates a microbial-enzyme model with 22 years of carbon cycling measurements at Harvard Forest. We find that increasing temperature reduces CUE but increases rB, and that two decades of soil warming increases the temperature sensitivities of CUE and rB. These temperature sensitivities, which are derived from decades-long field observations, contrast with values obtained from short-term laboratory experiments. We also show that long-term soil C flux and pool changes in response to warming are more dependent on the temperature sensitivity of CUE than that of rB. Using the inversion-derived parameters, we project that chronic soil warming at Harvard Forest over six decades will result in soil C gain of 1.0% on average (1st and 3rd quartiles: 3.0% loss and 10.5% gain) in the surface mineral horizon. Our results demonstrate that estimates of temperature sensitivity of microbial CUE and rB can be obtained and evaluated rigorously by integrating multidecadal datasets. This approach can potentially be applied in broader spatiotemporal scales to improve long-term projections of soil C feedbacks to climate warming.
Liao C, Luo Y, Tang X, Ma Z, Li B (2019) Effects of human population density on the pattern of terrestrial nature reserves in China. Global Ecology and Conservation 20: e00762.
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Read PublicationAn increasing number of studies showed that coverage of existing protected areas is not enough to protect biodiversity. However, to what extent and how human population density influence the geographical pattern of protected areas are not clear. Based on 2644 terrestrial nature reserves (NRs) in mainland China in 2015, correlation analysis showed that there was a significantly negative relationship between human density and area (R = −0.52, P < 0.001) and coverage of NRs (R = −0.21, P < 0.001), and a positive one between human density and density of NRs at county level (R = 0.64, P < 0.001) (all sample size n = 1171). These relationships could also be observed at provincial level. Counties with NRs had significantly lower human density (mean = 95 persons km−2) than those without (mean = 289 persons km−2) (P < 0.001, n = 31) across China. Both percentage of agricultural land and road density significantly and negatively correlated with area and coverage of NRs, and positively with human density and density of NRs at provincial level (all P < 0.01, n = 31). The relationships between human and NRs varied among 31 provinces, three conservation objectives of ecosystems, species and others, three hierarchical managements of national, provincial, and city-county levels, and two jurisdictional departments of forestry and non-forestry. But the general pattern of such relationships did not change. In addition, human density and density of NRs significantly positively, and area and coverage of NRs negatively correlated with density of IUCN red-list high plants and vertebrates excluding fishes at provincial level (all P < 0.05, n = 31). These results suggested that human density had substantial impacts on the geographical distribution of NRs when their sites were designated, elucidating the mechanism responsible for the low effectiveness of NRs in representing biodiversity.
Liu XJA, Finley BK, Mau RL, Schwartz E, Dijkstra P, Bowker MA, Hungate BA (2019) The soil priming effect: Consistent across ecosystems, elusive mechanisms. Soil Biology and Biochemistry 140: 107617.
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Read PublicationOrganic matter input to soils can accelerate the decomposition of native soil carbon (C), a process called the priming effect. Priming is ubiquitous and exhibits some consistent patterns, but a general explanation remains elusive, in part because of variation in the response across different ecosystems, and because of a diversity of proposed mechanisms, including microbial activation, stoichiometry, and community shifts. Here, we conducted five-week incubations of four soils (grassland, piñon-juniper, ponderosa pine, mixed conifer), varying the amount of substrate added (as 13C-glucose, either 350 or 1000 μg C g−1 week−1) and either with no added nitrogen (N), or with sufficient N (as NH4NO3) to bring the C-to-N ratio of the added substrate to 10. Using four different ecosystems enabled testing the generality of mechanisms underlying the priming effect. The responses of priming to the amount and C-to-N ratio of the added substrate were consistent across ecosystems: priming increased with the rate of substrate addition and declined when the C-to-N ratio of the substrate was reduced. However, structural equation models failed to confirm intermediate responses postulated to mediate the priming effect, including responses postulated to be mediated by stoichiometry and microbial activation. Specifically, priming was not clearly associated with changes in microbial biomass or turnover, nor with extracellular enzyme activities or the microbial C-to-N ratio. The strongest explanatory pathways in the structural equation models were the substrate, soil, and C-to-N ratio treatments themselves, with no intermediates, suggesting that either these measurements lacked sufficient sensitivity to reveal causal relationships, or the actual drivers for priming were not included in the ancillary measurements. While we observed consistent changes in priming caused by the amount and C-to-N ratio of the added substrate across a wide array of soils, our findings did not clearly conform to common models offered for the priming effect. Because priming is a residual flux involving diverse substrates of varying chemical composition, a simple and generalizable explanation of the phenomenon may be elusive.
Liu Y, Piao S, Gasser T, Ciais P, Yang H, Wang H, Keenan TF, Huang M, Wan S, Song J, Wang K, Janssens IA, Penuelas J, Huntingford C, Wang X, Altaf Arain M, Fang YJ, Michalak AM, Peng C, Poulter B, Schwalm C, Shi X, Tian H, Wei Y, Zeng N, Zhu Q, Wang T, (2019) Field-experiment constraints on the enhancement of the terrestrial carbon sink by CO2 fertilization. Nature Geoscience 12(10): 809-814.
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Read PublicationClarifying how increased atmospheric CO2 concentration (eCO2) contributes to accelerated land carbon sequestration remains important since this process is the largest negative feedback in the coupled carbon–climate system. Here, we constrain the sensitivity of the terrestrial carbon sink to eCO2 over the temperate Northern Hemisphere for the past five decades, using 12 terrestrial ecosystem models and data from seven CO2 enrichment experiments. This constraint uses the heuristic finding that the northern temperate carbon sink sensitivity to eCO2 is linearly related to the site-scale sensitivity across the models. The emerging data-constrained eCO2 sensitivity is 0.64 ± 0.28 PgC yr−1 per hundred ppm of eCO2. Extrapolating worldwide, this northern temperate sensitivity projects the global terrestrial carbon sink to increase by 3.5 ± 1.9 PgC yr−1 for an increase in CO2 of 100 ppm. This value suggests that CO2 fertilization alone explains most of the observed increase in global land carbon sink since the 1960s. More CO2 enrichment experiments, particularly in boreal, arctic and tropical ecosystems, are required to explain further the responsible processes.
Liu Y, Schwalm CR, Samules-Crow KE, Ogle K (2019) Ecological memory of daily carbon exchange across the globe and its importance in drylands. Ecology Letters 22(11): 1806-1816.
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Read PublicationHow do antecedent (past) conditions influence land-carbon dynamics after those conditions no longer persist? In particular, quantifying such memory effects associated with the influence of past environmental (exogenous) and biological (endogenous) conditions is crucial for understanding and predicting the carbon cycle. Here we show, using data from 42 eddy covariance sites across six major biomes, that ecological memory?decomposed into environmental and biological memory components?of daily net carbon exchange (NEE) is critical for understanding the land-carbon metabolism, especially in drylands for which memory explains ~ 32% of the variation in NEE. The strong environmental memory in drylands was primarily driven by short- and long-term moisture status. Moreover, the strength of environmental memory scales with increasing water stress. This universal scaling relationship, emerging within and among major biomes, suggests a potential adaptive response to water limitation. Our findings underscore the necessity of considering ecological memory in experiments, observations and modelling.
Marks JC (2019) Revisiting the fates of dead leaves that fall into streams. Annual Review of Ecology, Evolution, and Systematics 50(1): 547-568.
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Read PublicationAs terrestrial leaf litter decomposes in rivers, its constituent elements follow multiple pathways. Carbon leached as dissolved organic matter can be quickly taken up by microbes, then respired before it can be transferred to the macroscopic food web. Alternatively, this detrital carbon can be ingested and assimilated by aquatic invertebrates, so it is retained longer in the stream and transferred to higher trophic levels. Microbial growth on litter can affect invertebrates through three pathways, which are not mutually exclusive. First, microbes can facilitate invertebrate feeding, improving food quality by conditioning leaves and making them more palatable for invertebrates. Second, microbes can be prey for invertebrates. Third, microbes can compete with invertebrates for resources bound within litter and may produce compounds that retard carbon and nitrogen fluxes to invertebrates. As litter is broken down into smaller particles, there are many opportunities for its elements to reenter the stream food web. Here, I describe a conceptual framework for evaluating how traits of leaf litter will affect its fate in food webs and ecosystems that is useful for predicting how global change will alter carbon fluxes into and out of streams.
Moon TA, Overeem I, Druckenmiller M, Holland M, Huntington H, Kling G, Lovecraft AL, Miller G, Scambos T, Schaedel C, Schuur EAG, Trochim E, Wiese F, Williams D, Wong G (2019) The expanding footprint of rapid Arctic change. Earth's Future 7(3): 212-218.
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Read PublicationArctic land ice is melting, sea ice is decreasing, and permafrost is thawing. Changes in these Arctic elements are interconnected, and most interactions accelerate the rate of change. The changes affect infrastructure, economics, and cultures of people inside and outside of the Arctic, including in temperate and tropical regions, through sea level rise, worsening storm and hurricane impacts, and enhanced warming. Coastal communities worldwide are already experiencing more regular flooding, drinking water contamination, and coastal erosion. We describe and summarize the nature of change for Arctic permafrost, land ice, and sea ice, and its influences on lower latitudes, particularly the United States. We emphasize that impacts will worsen in the future unless individuals, businesses, communities, and policy makers proactively engage in mitigation and adaptation activities to reduce the effects of Arctic changes and safeguard people and society.
Morrissey EM, Mau RL, Hayer M, Liu XJA, Schwarz E, Dijkstra P, Koch BJ, Allen K, Blazewicz SJ, Hofmockel K, Pett-Ridge J, Hungate BA (2019) Evolutionary history constrains microbial traits across environmental variation. Nature Ecology & Evolution 3(7): 1064-1069.
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Read PublicationOrganisms influence ecosystems, from element cycling to disturbance regimes, to trophic interactions and to energy partitioning. Microorganisms are part of this influence, and understanding their ecology in nature requires studying the traits of these organisms quantitatively in their natural habitats—a challenging task, but one which new approaches now make possible. Here, we show that growth rate and carbon assimilation rate of soil microorganisms are influenced more by evolutionary history than by climate, even across a broad climatic gradient spanning major temperate life zones, from mixed conifer forest to high-desert grassland. Most of the explained variation (~50% to ~90%) in growth rate and carbon assimilation rate was attributable to differences among taxonomic groups, indicating a strong influence of evolutionary history, and taxonomic groupings were more predictive for organisms responding to resource addition. With added carbon and nitrogen substrates, differences among taxonomic groups explained approximately eightfold more variance in growth rate than did differences in ecosystem type. Taxon-specific growth and carbon assimilation rates were highly intercorrelated across the four ecosystems, constrained by the taxonomic identity of the organisms, such that plasticity driven by environment was limited across ecosystems varying in temperature, precipitation and dominant vegetation. Taken together, our results suggest that, similar to multicellular life, the traits of prokaryotes in their natural habitats are constrained by evolutionary history to a greater degree than environmental variation.
Natali SM, Watts JD, Rogers BM, Potter S, Ludwig SM, Selbmann AK, Sullivan PF, Abbott BW, Arndt KA, Birch L, Bjorkman MP, Bloom AA, Celis G, Christensen TR, Christensen CT, Commane R, Cooper EJ, Crill P, Czimczik C, Davydov S, Du J, Egan JE, Elberling B, Euskirchen ES, Friborg T, Genet H, Gockede M, Goodrich JP, Grogan P, Helbig M, Jafarov EE, Jastrow JD, Kalhori AAM, Kim Y, Kimball JS, Kuzbah L, Lara MJ, Larsen KS, Lee BY, Liu Z, Loranty MM, Lund M, Lupascu M, Madani N, Malhotra A, Matamala R, McFarland J, McGuire AD, Michelsen A, Minions C, Oechel WC, Olefeldt D, Parmentier FJW, Pirk N, Poulter B, Quinton W, Rezanezhad F, Risk D, Sachs T, Schaefer K, Schmidt NM, Schuur EAG, Semenchuk PR, Shaver G, Sonnentag O, Treat CC, Waldrop MP, Wang Y, Welker J, Wille C, Xu X, Zhang Z, Shuang O, Zona D (2019) Large loss of CO2 in winter observed across the northern permafrost region. Nature Climate Change 9(11):852-857.
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Read PublicationRecent warming in the Arctic, which has been amplified during the winter1–3, greatly enhances microbial decomposition of soil organic matter and subsequent release of carbon dioxide (CO2)4. However, the amount of CO2 released in winter is not known and has not been well represented by ecosystem models or empirically based estimates5,6. Here we synthesize regional in situ observations of CO2 flux from Arctic and boreal soils to assess current and future winter carbon losses from the northern permafrost domain. We estimate a contemporary loss of 1,662 TgC per year from the permafrost region during the winter season (October–April). This loss is greater than the average growing season carbon uptake for this region estimated from process models (−1,032 TgC per year). Extending model predictions to warmer conditions up to 2100 indicates that winter CO2 emissions will increase 17% under a moderate mitigation scenario—Representative Concentration Pathway 4.5—and 41% under business-as-usual emissions scenario—Representative Concentration Pathway 8.5. Our results provide a baseline for winter CO2 emissions from northern terrestrial regions and indicate that enhanced soil CO2 loss due to winter warming may offset growing season carbon uptake under future climatic conditions.
Nie J, Yi L, Xu H, Liu Z, Zeng Z, Dijkstra P, Koch GW, Hungate BA, Zhu B (2019) Leguminous cover crop Astragalus sinicus enhances grain yields and nitrogen use efficiency through increased tillering in an intensive double-cropping rice system in southern China. Agronomy 9(9).
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Read PublicationChinese milk vetch (Astragalus sinicus L., vetch), a leguminous winter cover crop, has been widely adopted by farmers in southern China to boost yield of the succeeding rice crop. However, the effects of vetch on rice grain yield and nitrogen (N) use efficiency have not yet been well studied in the intensive double-cropped rice cropping systems. To fill this gap, we conducted a three-year field experiment to evaluate the impacts of the vetch crop on yields and N use efficiency in the subsequent early and late rice seasons. With moderate N input (100 kg N ha&minus;1 for each rice crop), vetch cover significantly increased grain yields by 7.3&ndash;13.4% for early rice, by 8.2&ndash;10.4% for late rice, and by 8.6&ndash;11.5% for total annual rice production when compared with winter fallow. When rice crops received an N input of 200 kg N ha&minus;1, vetch cover increased grain yields by 5.9&ndash;18.4% for early rice, by 3.8&ndash;10.1% for late rice, and by 6.2&ndash;11.3% for annual rice production. Moreover, comparable grain yields (11.9 vs. 12.0 Mg ha&minus;1 for annual rice production) were observed between vetch cover with moderate N and fallow with added N fertilizer. Yield components analysis indicated that the increased tillering number was the main factor for the enhanced grain yields by vetch cover. Vetch cover with moderate and higher N input resulted in higher agronomic N use efficiency and applied N recovery efficiency compared with the fallow treatments. Here, our results showed that vetch as a winter cover crop can be combined with reduced N fertilizer input while maintaining high grain yields, thus gaining a more sustainable rice production system.
Ogle K, Peltier D, Fell M, Guo J, Kropp H, Barber J (2019) Should we be concerned about multiple comparisons in hierarchical Bayesian models?. Methods in Ecology and Evolution 10(4): 553-564.
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Read PublicationEcologists increasingly use hierarchical Bayesian (HB) models to estimate group-level parameters that vary by, for example, species, treatment level, habitat type or other factors. Group-level parameters may be compared to infer differences among levels. We would conclude a non-zero pairwise difference, separately, for each pair in the group, when the respective 95% credible interval excludes zero. Classical procedures suggest that the rejection procedure should be adjusted to control the family-wise error rate (FWER) for a family of differences. Adjustments for FWER have been considered unnecessary in HB models due to partial pooling whereby increased pooling strength - group-level parameters become more alike - could lead to decreased rejection rates (Type I error, FWER, or Power) and increased false acceptance rates (Type 2 error and its family-wise analogue). To address this, we conducted a simulation experiment with factors of sample size, group size, balance (missingness), overall mean and ratio of within- to between-group variances, resulting in 2016 factor-level combinations (scenarios), replicated 100 times, producing 201,600 pseudo datasets analysed in a Bayesian framework. We evaluated the results in the context of a new partial pooling index (PPI), which we show is also applicable to more complex model structures based on four real-data examples. Simulation results confirm intuition that rejection rates (false and true) decrease and false acceptance rates increase with increasing PPI or pooling strength (scenario-level R2 = 0.81-0.97). The relationship with PPI differed greatly for balanced versus unbalanced designs and was affected by group size, especially for family-wise errors. Critically, an HB model does not guarantee that the FWER will follow a set significance level (α); for example, even minor imbalance can lead to FWER > α for weak to moderate pooling. These results are confirmed by the real-data examples, suggesting that ecologists need to consider FWER when applying HB models, especially for large group sizes or incomplete datasets. Contrary to current thought, HB models are not immune to issues of multiplicity, and our proposed PPI offers a method for evaluating if a particular HB analysis is likely to produce FWER ≤ α (no adjustment or alternative solution required).
Papp K, Hungate BA, Schwartz E (2019) mRNA, rRNA and DNA quantitative stable isotope probing with H218O indicates use of old rRNA among soil Thaumarchaeota. Soil Biology and Biochemistry 130: 159-166.
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Read PublicationRNA is considered to be a short-lived molecule, indicative of cellular metabolic activity, whereas DNA is thought to turn over more slowly because living cells do not always grow and divide. To explore differences in the rates of synthesis of these nucleic acids, we used H218O quantitative stable isotope probing (qSIP) to measure the incorporation of 18O into 16S rRNA, the 16S rDNA, amoA mRNA and the amoA gene of soil Thaumarchaeota. Incorporation of 18O into the thaumarchaeal amoA mRNA pool was faster than into the 16S rRNA pool, suggesting that Thaumarchaea were metabolically active while using rRNA molecules that were likely synthetized prior to H218O addition. Assimilation rates of 18O into 16S rDNA and amoA genes were similar, which was expected because both genes are present in the same thaumarchaeal genome. The Thaumarchaea had significantly higher rRNA to rDNA ratios than bacteria, though the 18O isotopic signature of thaumarchaeal rRNA was lower than that of bacterial rRNA, further suggesting preservation of old non-labeled rRNA. Through qSIP of soil with H218O, we showed that 18O incorporation into thaumarchaeal nucleic acids was generally low, indicating slower turnover rates compared to bacteria, and potentially suggesting thaumarchaeal capability for preservation and efficient reuse of biomolecules.
Papp K, Hungate BA, Schwartz E (2019) Glucose triggers strong taxon-specific responses in microbial growth and activity: insights from DNA and RNA qSIP. Ecology.
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Read PublicationGrowth of soil microorganisms is often described as carbon limited, and adding labile carbon to soil often results in a transient and large increase in respiration. In contrast, soil microbial biomass changes little, suggesting that growth and respiration are decoupled in response to a carbon pulse. Alternatively, measuring bulk responses of the entire community (total respiration and biomass) could mask ecologically important variation among taxa in response to the added carbon. Here, we assessed taxon-specific variation in cellular growth (measured as DNA synthesis) and metabolic activity (measured as rRNA synthesis) following glucose addition to soil using quantitative stable isotope probing with H218O. We found that glucose addition altered rates of DNA and rRNA synthesis, but the effects were strongly taxon specific: glucose stimulated growth and rRNA transcription for some taxa, and suppressed these for others. These contrasting taxon-specific responses could explain the small and transient changes in total soil microbial biomass. Responses to glucose were not well predicted by a priori assignments of taxa into copiotrophic or oligotrophic categories. Across all taxa, rates of DNA and rRNA synthesis changed in parallel, indicating that growth and activity were coupled, and the degree of coupling was unaffected by glucose addition. This pattern argues against the idea that labile carbon addition causes a large reduction in metabolic growth efficiency; rather, the large pulse of respiration observed with labile substrate addition is more likely to be the result of rapid turnover of microbial biomass, possibly due to trophic interactions. Our results support a strong connection between rRNA synthesis and bacterial growth, and indicate that taxon-specific responses among soil bacteria can buffer responses at the scale of the whole community.
Pegoraro E, Mauritz M, Bracho R, Ebert C, Dijkstra P, Hungate BA, Konstantinidis KT, Luo Y, Schädel C, Tiedje JM, Zhou J, Shuur EAG (2019) Glucose addition increases the magnitude and decreases the age of soil respired carbon in a long-term permafrost incubation study. Soil Biology and Biochemistry 129: 201-211.
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Read PublicationHigher temperatures in northern latitudes will increase permafrost thaw and stimulate above- and belowground plant biomass growth in tundra ecosystems. Higher plant productivity increases the input of easily decomposable carbon (C) to soil, which can stimulate microbial activity and increase soil organic matter decomposition rates. This phenomenon, known as the priming effect, is particularly interesting in permafrost because an increase in C supply to deep, previously frozen soil may accelerate decomposition of C stored for hundreds to thousands of years. The sensitivity of old permafrost C to priming is not well known; most incubation studies last less than one year, and so focus on fast-cycling C pools. Furthermore, the age of respired soil C is rarely measured, even though old C may be vulnerable to labile C inputs. We incubated soil from a moist acidic tundra site in Eight Mile Lake, Alaska for 409 days at 15 °C. Soil from surface (0–25 cm), transition (45–55 cm), and permafrost (65–85 cm) layers were amended with three pulses of uniformly 13C-labeled glucose or cellulose every 152 days. Glucose addition resulted in positive priming in the permafrost layer 7 days after each substrate addition, eliciting a two-fold increase in cumulative soil C loss relative to unamended soils with consistent effects across all three pulses. In the transition and permafrost layers, glucose addition significantly decreased the age of soil-respired CO2-C with Δ14C values that were 115‰ higher. Previous field studies that measured the age of respired C in permafrost regions have attributed younger Δ14C ecosystem respiration values to higher plant contributions. However, the results from this study suggest that positive priming, due to an increase in fresh C supply to deeply thawed soil layers, can also explain the respiration of younger C observed at the ecosystem scale. We must consider priming effects to fully understand permafrost C dynamics, or we risk underestimating the contribution of soil C to ecosystem respiration.
Peltier DMP, Ogle K (2019) Legacies of La Niña: North American monsoon can rescue trees from winter drought. Global Change Biology 25(1): 121-133.
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Read PublicationWhile we often assume tree growth?climate relationships are time-invariant, impacts of climate phenomena such as the El Niño Southern Oscillation (ENSO) and the North American Monsoon (NAM) may challenge this assumption. To test this assumption, we grouped ring widths (1900-present) in three southwestern US conifers into La Niña periods (LNP) and other years (OY). The 4 years following each La Niña year are included in LNP, and despite 1-2 year growth declines, compensatory adjustments in tree growth responses result in essentially equal mean growth in LNP and OY, as average growth exceeds OY means 2-4 years after La Niña events. We found this arises because growth responses in the two periods are not interchangeable: Due to differences in growth?climate sensitivities and climatic memory, parameters representing LNP growth fail to predict OY growth and vice versa (decreases in R2 up to 0.63; lowest R2 = 0.06). Temporal relationships between growth and antecedent climate (memory) show warmer springs and longer growing seasons negatively impact growth following dry La Niña winters, but that NAM moisture can rescue trees after these events. Increased importance of monsoonal precipitation during LNP is key, as the largest La Niña-related precipitation deficits and monsoonal precipitation contributions both occur in the southern part of the region. Decreases in first order autocorrelation during LNP were largest in the heart of the monsoon region, reflecting both the greatest initial growth declines and the largest recovery. Understanding the unique climatic controls on growth in Southwest conifers requires consideration of both the influences and interactions of drought, ENSO, and NAM, each of which is likely to change with continued warming. While plasticity of growth sensitivity and memory has allowed relatively quick recovery in the tree-ring record, recent widespread mortality events suggest conditions may soon exceed the capacity for adjustment in current populations.
Plaza C, Pegoraro E, Bracho R, Celis G, Crummer KG, Hutchings JA, Hicks Pries CE, Maurtiz M, Natali SM, Salmon VG, Schaedel C, Webb EE, Schuur EAG (2019) Direct observation of permafrost degradation and rapid soil carbon loss in tundra. Nature Geoscience 12(8):627-631.
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Read PublicationEvidence suggests that 5–15% of the vast pool of soil carbon stored in northern permafrost ecosystems could be emitted as greenhouse gases by 2100 under the current path of global warming. However, direct measurements of changes in soil carbon remain scarce, largely because ground subsidence that occurs as the permafrost soils begin to thaw confounds the traditional quantification of carbon pools based on fixed depths or soil horizons. This issue is overcome when carbon is quantified in relation to a fixed ash content, which uses the relatively stable mineral component of soil as a metric for pool comparisons through time. We applied this approach to directly measure soil carbon pool changes over five years in experimentally warmed and ambient tundra ecosystems at a site in Alaska where permafrost is degrading due to climate change. We show a loss of soil carbon of 5.4% per year (95% confidence interval: 1.0, 9.5) across the site. Our results point to lateral hydrological export as a potential pathway for these surprisingly large losses. This research highlights the potential to make repeat soil carbon pool measurements at sentinel sites across the permafrost region, as this feedback to climate change may be occurring faster than previously thought.
Potter S, Solvik K, Erb A, Goetz SJ, Johnstone JF, Mack MC, Randerson JT, Roman MO, Schaaf CL, Turetsky MR, Ververbeke S, Walker XJ, Wang Z, Massey R, Rogers BM (2019) Climate change decreases the cooling effect from postfire albedo in boreal North America. Global Change Biology.
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Read PublicationFire is a primary disturbance in boreal forests and generates both positive and negative climate forcings. The influence of fire on surface albedo is a predominantly negative forcing in boreal forests, and one of the strongest overall, due to increased snow exposure in the winter and spring months. Albedo forcings are spatially and temporally heterogeneous and depend on a variety of factors related to soils, topography, climate, land cover/vegetation type, successional dynamics, time since fire, season, and fire severity. However, how these variables interact to influence albedo is not well understood, and quantifying these relationships and predicting postfire albedo becomes increasingly important as the climate changes and management frameworks evolve to consider climate impacts. Here we developed a MODIS-derived ?blue sky? albedo product and a novel machine learning modeling framework to predict fire-driven changes in albedo under historical and future climate scenarios across boreal North America. Converted to radiative forcing (RF), we estimated that fires generate an annual mean cooling of ?1.77 ± 1.35 W/m2 from albedo under historical climate conditions (1971?2000) integrated over 70 years postfire. Increasing postfire albedo along a south?north climatic gradient was offset by a nearly opposite gradient in solar insolation, such that large-scale spatial patterns in RF were minimal. Our models suggest that climate change will lead to decreases in mean annual postfire albedo, and hence a decreasing strength of the negative RF, a trend dominated by decreased snow cover in spring months. Considering the range of future climate scenarios and model uncertainties, we estimate that for fires burning in the current era (2016) the cooling effect from long-term postfire albedo will be reduced by 15%?28% due to climate change.
Prevey JS, Rixen C, Ruger N, Hoye TT, Bjorkman AD, Myers-Smith IH, Elmendorf SC, Ashton IW, Cannone N, Chisholm CL, Clark K, Cooper EJ, Elberling B, Fosaa AM, Henry GHR, Hollister RD, Jonsdottir IS, Klanderud K, Kopp CW, Levesque E, Mauritz M, Molau U, Natali SM, Oberbauer SF, Panchen ZA, Post E, Rumpf SB, Scmidt NM, Schuur E, Semenchuk PR, Smith JG, Suding KN, Totland O, Troxler T, Venn S, Wahren CH, Welker JM, Wipf S (2019) Warming shortens flowering seasons of tundra plant communities. Nature Ecology & Evolution 3(1): 45-52.
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Read PublicationAdvancing phenology is one of the most visible effects of climate change on plant communities, and has been especially pronounced in temperature-limited tundra ecosystems. However, phenological responses have been shown to differ greatly between species, with some species shifting phenology more than others. We analysed a database of 42,689 tundra plant phenological observations to show that warmer temperatures are leading to a contraction of community-level flowering seasons in tundra ecosystems due to a greater advancement in the flowering times of late-flowering species than early-flowering species. Shorter flowering seasons with a changing climate have the potential to alter trophic interactions in tundra ecosystems. Interestingly, these findings differ from those of warmer ecosystems, where early-flowering species have been found to be more sensitive to temperature change, suggesting that community-level phenological responses to warming can vary greatly between biomes.
Purcell AM, Dijkstra P, Finley B, Hayer M, Koch BJ, Mau RL, Morrissey E, Papp K, Schwartz E, Stone BW, Hungate BA (2019) Quantitative stable isotope probing with H218O to measure taxon-specific microbial growth. Methods of Soil Analysis 4.
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Read PublicationMicroorganisms in soil assimilate, transform, and mineralize soil C to support growth. There are an estimated 2.6 × 1029 microbial cells containing 26 Pg C in soils worldwide. Consequently, quantifying microbial growth in soil is critical for determining the degree to which microorganisms contribute to the global C cycle. Measuring taxon-specific microbial growth enables understanding of the contribution of microbial taxa to elemental transformations across ecosystems and their susceptibility to environmental perturbations. These measurements in soil have largely been lacking due to inadequate methods. Quantitative stable isotope probing (qSIP) with H218O is used to measure taxon-specific growth of microbial taxa in soil, an improvement compared with traditional stable isotope probing (SIP). In qSIP, DNA extracted from both a labeled (18O-enriched water) and an unlabeled treatment is separated into numerous density fractions by isopycnic centrifugation, and target genes are quantified and sequenced in each fraction. The taxon-specific DNA density shift and ultimately the isotopic composition (18O enrichment) is calculated for each taxon. Here we discuss methods and illustrate the procedure for quantifying microbial taxon-specific growth in soil with qSIP using heavy isotope enriched H218O.
Qiao Y, Wang J, Liang G, Du Z, Zhou J, Zhu C, Huang K, Zhou X, Luo Y, Yan L, Xia J (2019) Global variation of soil microbial carbon-use efficiency in relation to growth temperature and substrate supply. Nature Scientific Reports 9(1): 5621.
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Read PublicationSoil microbial carbon-use efficiency (CUE), which is defined as the ratio of growth over C uptake, is commonly assumed as a constant or estimated by a temperature-dependent function in current microbial-explicit soil carbon (C) models. The temperature-dependent function (i.e., CUE = CUE0 + m × (T − 20)) simulates the dynamic CUE based on the specific CUE at a given reference temperature (i.e., CUE0) and a temperature response coefficient (i.e., m). Here, based on 780 observations from 98 sites, we showed a divergent spatial distribution of the soil microbial CUE (0.5 ± 0.25; mean ± SD) at the global scale. Then, the key parameters CUE0 and m in the above equation were estimated as 0.475 and −0.016, respectively, based on the observations with the Markov chain Monte Carlo technique. We also found a strong dependence of microbial CUE on the type of C substrate. The multiple regression analysis showed that glucose influences the variation of measured CUE associated with the environmental factors. Overall, this study confirms the global divergence of soil microbial CUE and calls for the incorporation of C substrate beside temperature in estimating the microbial CUE in different biomes.
Quan Q, Tian D, Luo Y, Zhang F, Crowther TW, Zhu K, Chen HYH, Zhou Q, Niu S (2019) Water scaling of ecosystem carbon cycle feedback to climate warming. Science Advances 5(8): eaav1131.
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Read PublicationIt has been well established by field experiments that warming stimulates either net ecosystem carbon uptake or release, leading to negative or positive carbon cycle–climate change feedback, respectively. This variation in carbon-climate feedback has been partially attributed to water availability. However, it remains unclear under what conditions water availability enhances or weakens carbon-climate feedback or even changes its direction. Combining a field experiment with a global synthesis, we show that warming stimulates net carbon uptake (negative feedback) under wet conditions, but depresses it (positive feedback) under very dry conditions. This switch in carbon-climate feedback direction arises mainly from scaling effects of warming-induced decreases in soil water content on net ecosystem productivity. This water scaling of warming effects offers generalizable mechanisms not only to help explain varying magnitudes and directions of observed carbon-climate feedback but also to improve model prediction of ecosystem carbon dynamics in response to climate change.
Rademacher TT, Basler D, Eckes-Shepard AH, Fonti P, Friend AD, Le Moine J, Richardson AD (2019) Using direct phloem transport manipulation to advance understanding of carbon dynamics in forest trees. Frontiers in Forests and Global Change 2: 11.
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Read PublicationCarbon dynamics within trees are intrinsically important for physiological functioning, in particular growth and survival, as well as ecological interactions on multiple timescales. Thus, these internal dynamics play a key role in the global carbon cycle by determining the residence time of carbon in forests via allocation to different tissues and pools, such as leaves, wood, storage, and exudates. Despite the importance of tree internal carbon dynamics, our understanding of how carbon is used in trees, once assimilated, has major gaps. The primary tissue that transports carbon from sources to sinks within a tree is the phloem. Therefore, direct phloem transport manipulation techniques have the potential to improve understanding of numerous aspects of internal carbon dynamics. These include relationships between carbon assimilation, nonstructural carbon availability, respiration for growth and tissue maintenance, allocation to, and remobilization from, storage reserves, and long-term sequestration in lignified structural tissues. This review aims to: (1) introduce the topic of direct phloem transport manipulations, (2) describe the three most common methods of direct phloem transport manipulation and review their mechanisms, namely (i) girdling, (ii) compression and (iii) chilling; (3) summarize the known impacts of these manipulations on carbon dynamics and use in forest trees; (4) discuss potential collateral effects and compare the methods; and finally (5) highlight outstanding key questions that relate to tree carbon dynamics and use, and propose ways to address them using direct phloem transport manipulation.
Seyednasrollah B, Domec JC, Clark JS (2019) Spatiotemporal sensitivity of thermal stress for monitoring canopy hydrological stress in near real-time. Agricultural and Forest Meteorology 269-270: 220-230.
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Read PublicationMonitoring drought in real-time using minimal field data is a challenge for ecosystem management and conservation. Most methods require extensive data collection and in-situ calibration and accuracy is difficult to evaluate. Here, we demonstrated how the space-borne canopy “thermal stress”, defined as surface-air temperature difference, provides a reliable surrogate for drought-induced water stress in vegetation. Using physics-based relationships that accommodate uncertainties, we showed how changes in canopy water flux from ground-based measurements relate to both the surface energy balance and remotely-sensed thermal stress. Field measurements of evapotranspiration in the southeastern and northwestern US verify this approach based on sensitivity of evapotranspiration to thermal stress in a large range of atmospheric and climate conditions. We found that a 1 °C change in the thermal stress is comparable to 1–1.2 mm day−1 of evapotranspiration, depending on site and climate conditions. We quantified temporal and spatial sensitivity of evapotranspiration to the thermal stress and showed that it has the strongest relationship with evapotranspiration during warm and dry seasons, when monitoring drought is essential. Using only air and surface temperatures, we predicted the inter-annual anomaly in thermal stress across the contiguous United States over the course of 15 years and compared it with conventional drought indices. Among drought metrics that were considered in this study, the thermal stress had the highest correlation values. Our sensitivity results demonstrated that the thermal stress is a particularly strong indicator of water-use in warm seasons and regions. This simple metric can be used at varying time-scales to monitor surface evapotranspiration and drought in large spatial extents in near real-time.
Seyednasrollah B, Milliman T, Richardson AD (2019) Data extraction from digital repeat photography using xROI: An interactive framework to facilitate the process. ISPRS Journal of Photogrammetry and Remote Sensing 152: 132-144.
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Read PublicationDigital repeat photography and near-surface remote sensing have been used by environmental scientists to study environmental change for nearly a decade. However, a user-friendly, reliable, and robust platform to extract color-based statistics and time series from a large stack of images is still lacking. Here, we present an interactive open-source toolkit, called xROI, that facilitates the process of time series extraction and improves the quality of the final data. xROI provides a responsive environment for scientists to interactively (a) delineate regions of interest (ROI), (b) handle field of view (FOV) shifts, and (c) extract and export time series data characterizing color-based metrics. The software gives user the opportunity to adjust mask files or draw new masks, every time an FOV shift occurs. Utilizing xROI can significantly facilitate data extraction from digital repeat photography and enhance the accuracy and continuity of extracted data.
Song J, Wan S, Piao S, Knapp AK, Classen AT, Vicca S, Ciais P, Hovenden MJ, Leuzinger S, Beier C, Kardol P, Zia J, Liu Q, Ru J, Zhou Z, Luo Y, Guo D, Adam LJ, Zscheischler J, Dukes JS, Tang J, Chen J, Hofmockel KS, Kueppers LM, Rustad L, Liu L, Smith MD, Fatichi S, Shao P, Han H, Wang D, Lei L, Wang J, Zhang Q, Li X, Su F, Liu B, Yang F, Ma G, Li G, Liu Y, Liu Y, Yang Z, Zhang K, Miao Y, Hu M, Yan C, Zhang A, Zhong M, Hui Y, Li Y, Zheng M (2019) A meta-analysis of 1,119 manipulative experiments on terrestrial carbon-cycling responses to global change. Nature Ecology & Evolution 3(9): 1309-1320.
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Read PublicationDirect quantification of terrestrial biosphere responses to global change is crucial for projections of future climate change in Earth system models. Here, we synthesized ecosystem carbon-cycling data from 1,119 experiments performed over the past four decades concerning changes in temperature, precipitation, CO2 and nitrogen across major terrestrial vegetation types of the world. Most experiments manipulated single rather than multiple global change drivers in temperate ecosystems of the USA, Europe and China. The magnitudes of warming and elevated CO2 treatments were consistent with the ranges of future projections, whereas those of precipitation changes and nitrogen inputs often exceeded the projected ranges. Increases in global change drivers consistently accelerated, but decreased precipitation slowed down carbon-cycle processes. Nonlinear (including synergistic and antagonistic) effects among global change drivers were rare. Belowground carbon allocation responded negatively to increased precipitation and nitrogen addition and positively to decreased precipitation and elevated CO2. The sensitivities of carbon variables to multiple global change drivers depended on the background climate and ecosystem condition, suggesting that Earth system models should be evaluated using site-specific conditions for best uses of this large dataset. Together, this synthesis underscores an urgent need to explore the interactions among multiple global change drivers in underrepresented regions such as semi-arid ecosystems, forests in the tropics and subtropics, and Arctic tundra when forecasting future terrestrial carbon-climate feedback.
Still C, Powell R, Aubrecht D, Kim Y, Helliker B, Roberts D, Richardson AD, Goulden M (2019) Thermal imaging in plant and ecosystem ecology: Applications and challenges. Ecosphere 10(6): e02768.
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Read PublicationTemperature is a primary environmental control on ecological systems and processes at a range of spatial and temporal scales. The surface temperature of organisms is often more relevant for ecological processes than air temperature, which is much more commonly measured. Surface temperature influences?and is influenced by?a range of biological, physical, and chemical processes, providing a unique view of temperature effects on ecosystem function. Furthermore, surface temperatures vary markedly over a range of temporal and spatial scales and may diverge from air temperature by 40°C or more. Surface temperature measurements have been challenging due to sensor and computational limitations but are now feasible at high spatial and temporal resolutions using thermal imaging. Thus, significant advances in our understanding of plant and ecosystem thermal regimes and their functional consequences are now possible. Thermal measurements may be used to address many ecological questions, such as the thermal controls on plant and ecosystem metabolism and the impact of heat waves and drought. Further advances in this area will require interdisciplinary collaborations among practitioners in fields ranging from physiology to ecosystem ecology to remote sensing and geospatial analysis. In this overview, we demonstrate the feasibility, utility, and potential of thermal imaging for measuring vegetation surface temperatures across a range of scales and from measurement, analysis, and synthesis perspectives.
Stuble KL, Ma S, Liang J, Luo Y, Classen AT, Souza L (2019) Long-term impacts of warming drive decomposition and accelerate the turnover of labile, not recalcitrant, carbon. Ecosphere 10(5): e02715.
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Read PublicationWarming is altering the way soils function in ecosystems both directly by changing microbial physiology and indirectly by causing shifts in microbial community composition. Some of these warming-driven changes are short term, but others may persist over time. Here, we took advantage of a long-term (14 yr) warming experiment in a tallgrass prairie to tease apart the influence of short- and long-term warming on litter decomposition. We collected soils originating from warmed and control plots and incubated them with a common litter substrate in a reciprocal design under elevated and ambient growth chamber temperatures. Litter decomposition was 40% higher in soils that were warmed in the field for 14 yr (long-term warming) relative to soils derived from ambient plots. Short-term warming in the laboratory had less of an impact on decomposition-decomposition increased by 12% under laboratory warming. Using a two-pool soil carbon model to explore how different carbon pools may be responding, we found that long-term warming accelerated the turnover of labile, not recalcitrant, carbon in these prairie soils-a result that is likely due to shifts in soil community activity/composition. Taken together, our results offer experimental evidence that warming-induced changes in the soil community that occur over 14 yr of warming have long-lasting effects on carbon turnover.
Terrer C, Jackson RB, Prentice IC, Kennan TF, Kaiser C, Vicca S, Fisher JB, Reich PB, Stocker BD, Hungate BA, Penuelas J, McCallum I, Soudzilovskaia NA, Cernusak LA, Talhelm AF, Van Sundert K, Piao S, Nwton PCD, Hovenden MJ, Blumenthal DM, Liu YY, Muller C, Winter K, Field CB, Viechtbauer W, Van Lissa CJ, Hoosbeek MR, Watanabe M, Koike T, Leshyk VO, Polley HW, Franklin O (2019) Nitrogen and phosphorus constrain the CO2 fertilization of global plant biomass. Nature Climate Change 9(9): 684-689.
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Read PublicationElevated CO2 (eCO2) experiments provide critical information to quantify the effects of rising CO2 on vegetation1–6. Many eCO2 experiments suggest that nutrient limitations modulate the local magnitude of the eCO2 effect on plant biomass1,3,5, but the global extent of these limitations has not been empirically quantified, complicating projections of the capacity of plants to take up CO27,8. Here, we present a data-driven global quantification of the eCO2 effect on biomass based on 138 eCO2 experiments. The strength of CO2 fertilization is primarily driven by nitrogen (N) in ~65% of global vegetation and by phosphorus (P) in ~25% of global vegetation, with N- or P-limitation modulated by mycorrhizal association. Our approach suggests that CO2 levels expected by 2100 can potentially enhance plant biomass by 12 ± 3% above current values, equivalent to 59 ± 13 PgC. The global-scale response to eCO2 we derive from experiments is similar to past changes in greenness9 and biomass10 with rising CO2, suggesting that CO2 will continue to stimulate plant biomass in the future despite the constraining effect of soil nutrients. Our research reconciles conflicting evidence on CO2 fertilization across scales and provides an empirical estimate of the biomass sensitivity to eCO2 that may help to constrain climate projections.
Thomas, Scott C Tamadonfar, Kevin O Symour, Cale O Lai, Dengxun Dodsworth, Jeremy A Murugapiran, Senthil K Eloe-Fadrosh, Emiley A Dijkstra, Paul Hedlund, Brian P (2019) Position-specific metabolic probing and metagenomics of microbial communities reveal conserved central carbon metabolic network activities at high temperatures. Frontiers in Microbiology 10.
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Read PublicationTemperature is a primary driver of microbial community composition and taxonomic diversity; however, it is unclear to what extent temperature affects characteristics of central carbon metabolic pathways (CCMPs). In this study, 16S rRNA gene amplicon and metagenome sequencing were combined with 13C-labeled metabolite probing of the CCMPs to assess community carbon metabolism along a temperature gradient (60-95 °C) in Great Boiling Spring, NV. 16S rRNA gene amplicon diversity was inversely proportional to temperature, and Archaea were dominant at higher temperatures. Metagenomes spanning the temperature gradient hosted abundant CCMPs genes and many individual metagenome-assembled genomes had complete pathways. In contrast, genes encoding cellulosomes and some others involved in plant matter degradation and most for photosynthesis and were absent at higher temperatures. In situ 13C-CO2 production from labeled isotopomer pairs of glucose, pyruvate, and acetate suggested lower relative oxidative pentose phosphate pathway activity and/or fermentation at 60°C, and a stable or decreased maintenance energy demand at higher temperatures. Catabolism of 13C-labeled citrate, succinate, L-alanine, L-serine, and L-cysteine was observed at 85°C, demonstrating broad heterotrophic activity. Together, these results suggest that temperature-driven losses in biodiversity in geothermal systems may not alter CCMP function or maintenance energy demands at a community level.
Turetsky MR, Abbott BW, Jones MC, Walter Anthony K, Olefeldt D, Schuur EAG, Koven C, McGuire AD, Grosse G, Kuhry P, Hugelius G, Lawrence DM, Gibson C, Sannel ABK (2019) Permafrost collapse is accelerating carbon release. Nature 569.
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Read PublicationWalker AP, De Kauwe MG, Medlyn BE, Zaehle S, Iversen CM, Asao S, Guenet B, Harper A, Hickler T, Hungate BA, Jain AK, Luo Y, Lu X, Luus K, Megonigal JP, Oren R, Ryan E, Shu S, Talhelm A, Wang YP, Warren JM, Werner C, Xia J, Yang B, Zak DR, Norby RJ (2019) Decadal biomass increment in early secondary succession woody ecosystems is increased by CO2 enrichment. Nature Communications 10(1): 454.
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Read PublicationIncreasing atmospheric CO2 stimulates photosynthesis which can increase net primary production (NPP), but at longer timescales may not necessarily increase plant biomass. Here we analyse the four decade-long CO2-enrichment experiments in woody ecosystems that measured total NPP and biomass. CO2 enrichment increased biomass increment by 1.05 ± 0.26 kg C m−2 over a full decade, a 29.1 ± 11.7% stimulation of biomass gain in these early-secondary-succession temperate ecosystems. This response is predictable by combining the CO2 response of NPP (0.16 ± 0.03 kg C m−2 y−1) and the CO2-independent, linear slope between biomass increment and cumulative NPP (0.55 ± 0.17). An ensemble of terrestrial ecosystem models fail to predict both terms correctly. Allocation to wood was a driver of across-site, and across-model, response variability and together with CO2-independence of biomass retention highlights the value of understanding drivers of wood allocation under ambient conditions to correctly interpret and predict CO2 responses.
Walker XJ, Baltzer JL, Cumming SG, Day NJ, Ebert C, Goetz S, Johnstone JF, Potter S, Rogers BM, Schuur EAG, Turetsky MR, Mack MC (2019) Increasing wildfires threaten historic carbon sink of boreal forest soils. Nature 572(7770): 520-523.
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Read PublicationBoreal forest fires emit large amounts of carbon into the atmosphere primarily through the combustion of soil organic matter1–3. During each fire, a portion of this soil beneath the burned layer can escape combustion, leading to a net accumulation of carbon in forests over multiple fire events4. Climate warming and drying has led to more severe and frequent forest fires5–7, which threaten to shift the carbon balance of the boreal ecosystem from net accumulation to net loss1, resulting in a positive climate feedback8. This feedback will occur if organic-soil carbon that escaped burning in previous fires, termed ‘legacy carbon’, combusts. Here we use soil radiocarbon dating to quantitatively assess legacy carbon loss in the 2014 wildfires in the Northwest Territories of Canada2. We found no evidence for the combustion of legacy carbon in forests that were older than the historic fire-return interval of northwestern boreal forests9. In forests that were in dry landscapes and less than 60 years old at the time of the fire, legacy carbon that had escaped burning in the previous fire cycle was combusted. We estimate that 0.34 million hectares of young forests (<60 years) that burned in the 2014 fires could have experienced legacy carbon combustion. This implies a shift to a domain of carbon cycling in which these forests become a net source—instead of a sink—of carbon to the atmosphere over consecutive fires. As boreal wildfires continue to increase in size, frequency and intensity7, the area of young forests that experience legacy carbon combustion will probably increase and have a key role in shifting the boreal carbon balance.
Wang H, Shen M, Hui D, Chen J, Sun G, Wang X, Lu C, Sheng J, Chen L, Luo Y, Zheng J, Zhang Y (2019) Straw incorporation influences soil organic carbon sequestration, greenhouse gas emission, and crop yields in a Chinese rice (Oryza sativa L.) –wheat (Triticum aestivum L.) cropping system. Soil and Tillage Research 195:104377.
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Read PublicationCrop straw management plays important roles in sustainable agriculture and environmental protection. Straw incorporation has multiple influences on soil organic carbon (SOC) sequestration, greenhouse gas (GHG) emissions, and crop yields, but these influences have rarely been studied simultaneously in a single cropping system. This study was conducted to examine the influence of long-term straw incorporation on the SOC sequestration rate, methane (CH4) and nitrous oxide (N2O) emissions and crop yields in a Chinese rice (Oryza sativa L.) –wheat (Triticum aestivum L.) cropping system in Hydragric Anthrosols under a subtropical monsoon climate. Four straw incorporation treatments were applied: wheat straw incorporation only (WS), rice straw incorporation only (RS), both wheat and rice straw incorporation (WSRS), and no straw incorporation (as a control). The SOC sequestration rate was estimated from the changes in SOC stock in the topsoil (0–20 cm) from 2007 to 2016. The emissions of CH4 and N2O were measured every 7 d when possible using a static chamber method from the 2013 rice season to the 2016 wheat season. Our results showed that the straw incorporation treatments significantly influenced the seasonal CH4 and N2O emissions and rice yield but had no influence on wheat yield. Straw incorporation significantly increased the annual topsoil SOC sequestration rate by 0.24–0.43 t C ha−1 yr−1 and the annual CH4 and N2O emissions by 44–138 kg CH4-C ha−1 yr−1 and 0.68–1.49 kg N2O-N ha−1 yr−1, respectively. Relative to the RS treatment, the WS and WSRS treatments significantly increased annual CH4 emissions by 38% and 61%, respectively. Relative to the RS treatment, the WSRS treatment significantly increased the annual N2O emissions, by 35%. The average annual yields were significantly higher in the WSRS (16.8 t ha−1 yr−1) and RS (16.7 t ha−1 yr−1) treatments than in the WS (15.7 t ha−1 yr−1) and control (15.2 t ha−1 yr−1) treatments. Across the three rotation cycles, the annual net global warming potential and greenhouse gas intensity were similar between the control and RS treatments but were significantly lower in these treatments than in the WSRS and WS treatments. These findings suggest that the RS treatment can simultaneously increase crop yields and environmental sustainability in rice–wheat cropping systems.
Wang J, Xia J, Zhou X, Huang K, Zhou J, Huang Y, Jiang L, Xu X, Liang J, Wang YP, Cheng X, Luo Y (2019) Evaluating the simulated mean soil carbon transit times by Earth system models using observations. Biogeosciences 16: 917-926.
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Read PublicationOne known bias in current Earth system models (ESMs) is the underestimation of global mean soil carbon (C) transit time (<span class="inline-formula"><i>τ</i><sub>soil</sub></span>), which quantifies the age of the C atoms at the time they leave the soil. However, it remains unclear where such underestimations are located globally. Here, we constructed a global database of measured <span class="inline-formula"><i>τ</i><sub>soil</sub></span> across 187 sites to evaluate results from 12 ESMs. The observations showed that the estimated <span class="inline-formula"><i>τ</i><sub>soil</sub></span> was dramatically shorter from the soil incubation studies in the laboratory environment (median <span class="inline-formula">=</span> 4 years; interquartile range <span class="inline-formula">=</span> 1 to 25 years) than that derived from field in situ measurements (31; 5 to 84 years) with shifts in stable isotopic C (<span class="inline-formula"><sup>13</sup>C</span>) or the <i>stock-over-flux</i> approach. In comparison with the field observations, the multi-model ensemble simulated a shorter median (19 years) and a smaller spatial variation (6 to 29 years) of <span class="inline-formula"><i>τ</i><sub>soil</sub></span> across the same site locations. We then found a significant and negative linear correlation between the in situ measured <span class="inline-formula"><i>τ</i><sub>soil</sub></span> and mean annual air temperature. The underestimations of modeled <span class="inline-formula"><i>τ</i><sub>soil</sub></span> are mainly located in cold and dry biomes, especially tundra and desert. Furthermore, we showed that one ESM (i.e., CESM) has improved its <span class="inline-formula"><i>τ</i><sub>soil</sub></span> estimate by incorporation of the soil vertical profile. These findings indicate that the spatial variation of <span class="inline-formula"><i>τ</i><sub>soil</sub></span> is a useful benchmark for ESMs, and we recommend more observations and modeling efforts on soil C dynamics in regions limited by temperature and moisture.
Williams CB, Anfodillo T, Crivellaro A, Lazzarin M, Dawson TE, Koch GW (2019) Axial variation of xylem conduits in the Earth’s tallest trees. Trees.
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Read PublicationHydraulic limitations to tree height can be mitigated by widening the conducting elements toward a tree’s base. However, size limits of tracheid and vessel dimensions may constrain this compensation mechanism as the water transport pathway elongates. Moreover, variation in conduit size is poorly described in tall trees even though their long transport paths have high potential for hydraulic resistance. Here, we evaluated whether axial variation in conduit diameter was uniquely structured, or matched theoretical predictions in <em class="EmphasisTypeItalic ">Sequoia sempervirens</em>, <em class="EmphasisTypeItalic ">Sequoiadendron giganteum,</em> and <em class="EmphasisTypeItalic ">Eucalyptus regnans</em> that were 86–105 m tall and exceeded 85% of the maximum height for each species. Across <em class="EmphasisTypeItalic ">Sequoia</em> and <em class="EmphasisTypeItalic ">Sequoiadendron</em>, tree top tracheids maintained constant width, whereas tree base tracheids in the outermost ring were 20% wider in taller trees, suggesting maintenance of basipetal conduit widening with height growth. In all trees, the observed widening decreased at a rate per unit path length that fitted well to a power function with an exponent consistent with hydraulic compensation. However, below about 60 m from the tree tops, conduit diameters approached an asymptote beneath the power function, indicating a limit to maximum conduit size. Quantifying the distribution of base-to-top hydraulic resistance suggested that the minimal hydraulic benefit gained with increasingly wider conduits near the tree base may trade off with other factors such as maintaining mechanical strength or reducing fluid volume. We summarize these results into an anatomical model of height growth that includes limits to axial variation in conduit diameter and is supported by many physiological and anatomical observations.
Wright CS, Evans AM, Grove S, Haubensak KA (2019) Pile age and burn season influence fuelbed properties, combustion dynamics, fuel consumption, and charcoal formation when burning hand piles. Forest Ecology and Management 439: 146-158.
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Read PublicationPiling and burning is widely used to dispose of unmerchantable debris resulting from thinning in forests throughout the western United States. Quite often more piles are created than are burned in a given year, however, causing piles to persist, accumulate, and age on the landscape. The effects of burning piles of increasing age has not been studied. We examined the effects of time since construction (i.e., pile age, in roughly six month increments for two years) and burn season (fall and spring) on fuelbed properties, combustion dynamics, fuel consumption, and charcoal formation for hand-constructed piles in thinned ponderosa pine-dominated sites in New Mexico (n = 50 piles) and Washington (n = 49 piles). Piles compacted over time similarly for both study sites, losing approximately 15% of their height annually for the first two years following piling. Peak flame height decreased and the duration of flaming combustion increased with increasing pile age for both burn seasons in New Mexico, yet depended on burn season in Washington. Increasing fuel moisture and compaction reduced peak flame height and increased flaming duration modestly for both sites. Peak flame height was reduced 6–7 cm and flaming duration increased 0.9–2.3 min for every percentage increase in small fuel moisture. Similarly, peak flame height was reduced 4–5 cm and flaming duration increased 0.6–0.8 min for every percentage reduction in pile height. Fuel consumption was high, averaging 90% in New Mexico and 95% in Washington. Fuel consumption patterns differed between locations, however; fuel consumption decreased with age and was slightly higher for spring than fall burns in New Mexico, whereas, neither pile age nor burn season affected fuel consumption in Washington. Charcoal formation as a fraction of pre-burn pile weight averaged 2.8% in New Mexico and 1.2% in Washington, and was not affected by pile age or burn season. Fuel consumption and charcoal production were unaffected by fuel moisture or compaction levels at either site. Findings from this study will inform fuel and fire managers about the potential effects on fire behavior, fuel consumption, and charcoal formation of burning piles of different age in different seasons under different environmental conditions.
Wu Z, Hugelius G, Luo Y, Smith B, Xia J, Fensholt R, Lehsten V, Ahlstrom A (2019) Approaching the potential of model-data comparisons of global land carbon storage. Scientific Reports 9(1): 3367.
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Read PublicationCarbon storage dynamics in vegetation and soil are determined by the balance of carbon influx and turnover. Estimates of these opposing fluxes differ markedly among different empirical datasets and models leading to uncertainty and divergent trends. To trace the origin of such discrepancies through time and across major biomes and climatic regions, we used a model-data fusion framework. The framework emulates carbon cycling and its component processes in a global dynamic ecosystem model, LPJ-GUESS, and preserves the model-simulated pools and fluxes in space and time. Thus, it allows us to replace simulated carbon influx and turnover with estimates derived from empirical data, bringing together the strength of the model in representing processes, with the richness of observational data informing the estimations. The resulting vegetation and soil carbon storage and global land carbon fluxes were compared to independent empirical datasets. Results show model-data agreement comparable to, or even better than, the agreement between independent empirical datasets. This suggests that only marginal improvement in land carbon cycle simulations can be gained from comparisons of models with current-generation datasets on vegetation and soil carbon. Consequently, we recommend that model skill should be assessed relative to reference data uncertainty in future model evaluation studies.
Xu Y, Ida Y, Huang H, Shi Z, Franklin SB, Luo Y, Bao D, Qiao X, Jiang M (2019) Linkages between tree architectural designs and life-history strategies in a subtropical montane moist forest. Forest Ecology and Management 438(15: 1-9.
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Read PublicationTree architecture is crucial to maximizing light capture, determined by carbon allocation of individual trees, and consequently characterizes species-specific growth strategies. Its variation and associated life-history strategies have been examined in tropical and temperate forests, but not in subtropical forests. Moreover, a similar architectural pattern was found using a hierarchical Bayesian model in a tropical forest, which differed from most of previous studies. Here, we employed a hierarchical Bayesian model to examine tree architecture differences and associations with adult stature and light requirement among 59 subtropical co-occurring species. Architectural variations among tree species with different seed dispersal and leaf phenology types were analyzed. Most species showed similar architecture in the height of the lowest foliage-tree height relationships (F-H) and the long side of crown- tree height relationships (W1-H), but some species showed interspecific variations in tree height-stem diameter relationships (H-D) among the 59 co-occurring species in the subtropical montane forest. Trees developed deeper and larger crowns at mid-elevation compared to the tropical and temperate forests. Parameters of H-D relationship differed in leaf phenology and dispersal types, and intercepts of F-H relationship and W1-H relationship differed in leaf phenology. Large-statured species had more slender stems, and shallower and narrower crowns at small sizes, but similar crowns at large sizes. Light-demanding species showed weak correlations between architectural variables and light requirement but exhibited wide crowns at the intermediate sizes. In general, size-dependent architectural differentiation was driven mainly by adult stature and light requirement in subtropical forest. Coexistence species showed different life-history strategies in light capture, which may help provide options in forest thinning and harvesting in subtropical forest. Species-specific tree architectural models of 59 co-occurring species represent three-dimensional (3D) structure of this subtropical forest accurately, but also support for future terrestrial laser scanning (TLS) data analysis.
Yan Q, Dong F, Yang F, Lu J, Li F, Zhang J, Dong J, Li J (2019) Improved yield and water storage of the wheat-maize rotation system due to double-blank row mulching during the wheat stage. Agricultural Water Management 213: 903-912.
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Read PublicationMulching techniques have been widely used in dryland regions in northern China. It is necessary to develop water-saving cultivation techniques in irrigation regions in northern China to relieve water scarcity. Planting and mulching on separate rows has been widely used to improve wheat yield and involves a pattern of a double row of planting and a blank row of mulching. However, whether the mulching pattern during the wheat season can be applied to the wheat-maize system to increase the yield of both crops and to reduce the use of irrigation water remains unclear. Three mulching practices (conventional planting (CP), conventional planting with mulching (CPM) and double-blank planting with mulching (DPM)) during the wheat season were conducted to verify the potential roles of DPM in increasing wheat and maize yields, improving soil temperature and enhancing water storage under the DPM practice. The results show that the DPM practice significantly increased the efficiency spike number, aboveground biomass and grain yield (7.8% higher than CP and 11.3% higher than CPM) of wheat. The heat conservation effect of the DPM practice was stronger in the early stage of growth and was more effective in minimizing fluctuations in soil temperature in the wheat season compared with CPM. The development and yield of maize that was sowed in the mulching lines of DPM were less improved, although the amount of aboveground biomass at the maturity stage was higher. Additionally, the soil temperature of the maize season under DPM showed a narrowing trend of changes during the early stage with slight effects in the middle stage and a resumption of heat conservation in the late stage. Compared with CP, both mulching patterns decreased soil evaporation during the two crops’ seasons by an average 5.3% in CPM and 7.8% in DPM, which is particularly evident when the crops’ leaf area index was low. Therefore, the DPM pattern could more effectively optimize soil temperature and water storage. Furthermore, this pattern may have positive effects on the yields of winter wheat and on reducing the soil water requirement of the maize season.
Yang S, Zheng Q, Yuan M, Shi Z, Chiariello NR, Docherty KM, Dong S, Field CB, Gu Y, Gutknecht J, Hungate BA, Le Roux X, Ma X, Niboyet A, Yuan T, Zhou J, Yang Y (2019) Long-term elevated CO2 shifts composition of soil microbial communities in a Californian annual grassland, reducing growth and N utilization potentials. Science of The Total Environment 652: 1474-1481.
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Read PublicationThe continuously increasing concentration of atmospheric CO2 has considerably altered ecosystem functioning. However, few studies have examined the long-term (i.e. over a decade) effect of elevated CO2 on soil microbial communities. Using 16S rRNA gene amplicons and a GeoChip microarray, we investigated soil microbial communities from a Californian annual grassland after 14 years of experimentally elevated CO2 (275 ppm higher than ambient). Both taxonomic and functional gene compositions of the soil microbial community were modified by elevated CO2. There was decrease in relative abundance for taxa with higher ribosomal RNA operon (rrn) copy number under elevated CO2, which is a functional trait that responds positively to resource availability in culture. In contrast, taxa with lower rrn copy number were increased by elevated CO2. As a consequence, the abundance-weighted average rrn copy number of significantly changed OTUs declined from 2.27 at ambient CO2 to 2.01 at elevated CO2. The nitrogen (N) fixation gene nifH and the ammonium-oxidizing gene amoA significantly decreased under elevated CO2 by 12.6% and 6.1%, respectively. Concomitantly, nitrifying enzyme activity decreased by 48.3% under elevated CO2, albeit this change was not significant. There was also a substantial but insignificant decrease in available soil N, with both nitrate (NO3−) (−27.4%) and ammonium (NH4+) (−15.4%) declining. Further, a large number of microbial genes related to carbon (C) degradation were also affected by elevated CO2, whereas those related to C fixation remained largely unchanged. The overall changes in microbial communities and soil N pools induced by long-term elevated CO2 suggest constrained microbial N decomposition, thereby slowing the potential maximum growth rate of the microbial community.
Yang W, Jeelani N, Zhu Z, Luo Y, Cheng X, An S (2019) Alterations in soil bacterial community in relation to Spartina alterniflora Loisel. invasion chronosequence in the eastern Chinese coastal wetlands. Applied Soil Ecology 135: 38-43.
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Read PublicationIn order to better understand the variations in soil bacterial community and associated drivers following plant invasion, we investigated changes in soil bacterial community along with 9-, 13-, 20- and 23-year-old Spartina alterniflora Loisel. (SA) invasion in comparison with bare flat (BF) in the eastern Chinese coastal wetlands, based on analyses of quantitative polymerase chain reaction (qPCR) and Illumina MiSeq DNA sequencing of 16S rRNA gene. The SA invasion significantly elevated soil bacterial abundance and diversity relative to BF, with the highest levels in 9-year-old SA soil, which gradually decreased with SA invasion from 9 to 23 years. The abundance of copiotrophic Proteobacteria, β-proteobacteria, and Bacteroidetes generally diminished along with SA invasion chronosequence. While, changes in abundance of oligotrophic Chloroflexi, Acidobacteria, Nitrospirae and Planctomycetes exhibited opposite trends. Our data suggest that soil nutrient substrates, and physiochemical properties (soil pH and/or moisture) primarily drive the shifts in soil bacterial abundance, diversity, and community composition along with SA invasion chronosequence in the costal wetlands of eastern China. Overall, soil bacterial abundance and diversity peaked in 9-year-old SA community, with soil bacterial community composition changing from copiotrophic to oligotrophic groups along with SA invasion chronosequence.
Yang W, Zhang D, Cai X, Xia L, Luo Y, Cheng X, An S (2019) Significant alterations in soil fungal communities along a chronosequence of Spartina alterniflora invasion in a Chinese Yellow Sea coastal wetland. Science of the Total Environment 693: 133548.
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Read PublicationPlant invasion typically alters the microbial communities of soils, which affects ecosystem carbon (C) and nitrogen (N) cycles. The responses of the soil fungal communities to plant invasion along its chronosequence remain poorly understood. For this study, we investigated variations in soil fungal communities through Illumina MiSeq sequencing analyses of the fungal internal transcribed spacer (ITS) region, and quantitative polymerase chain reaction (qPCR), along a chronosequence (i.e., 9-, 13-, 20- and 23-year-old) of invasive Spartina alterniflora. We compared these variations with those of bare flat in a Chinese Yellow Sea coastal wetland. Our results highlighted that the abundance of soil fungi, the number of operational taxonomic units (OTUs), species richness, and Shannon diversity indices for soil fungal communities were highest in 9-year-old S. alterniflora soil, which gradually declined along the invasion chronosequence. The relative abundance of copiotrophic Basidiomycota revealed significant decreasing trend, while the relative abundance of oligotrophic Ascomycota gradually increased along the S. alterniflora invasion chronosequence. The relative abundance of soil saprotrophic fungi (e.g., undefined saprotrophs) was gradually reduced while symbiotic fungi (e.g., ectomycorrhizal fungi) and pathotrophic fungi (e.g., plant and animal pathogens) progressively increased along the S. alterniflora invasion chronosequence. Our results suggested that S. alterniflora invasion significantly altered soil fungal abundance and diversity, community composition, trophic modes, and functional groups along a chronosequence, via substantially reduced soil litter inputs, and gradually decreased soil pH, moisture, and soil nutrient substrates along the invasion chronosequence, from 9 to 23 years. These changes in soil fungal communities, particularly their trophic modes and functional groups along the S. alterniflora invasion chronosequence could well impact the decomposition and accumulation of soil C and N, while potentially altering ecosystem C and N sinks in a Chinese Yellow Sea coastal wetland.
Zhang M, Cheng X, Geng Q, Shi Z, Luo Y, Xu X (2019) Leaf litter traits predominantly control litter decomposition in streams worldwide. Global Ecology and Biogeography 28(10):1469-1486.
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Read PublicationAim Leaf litter decomposition in freshwater ecosystems is a vital process linking ecosystem nutrient cycling, energy transfer and trophic interactions. In comparison to terrestrial ecosystems, in which researchers find that litter traits predominantly regulate litter decomposition worldwide, the dominant factors controlling its decomposition in aquatic ecosystems are still debated, with global patterns not well documented. Here, we aimed to explore general patterns and key drivers (e.g., litter traits, climate and water characteristics) of leaf litter decomposition in streams worldwide. Location Global. Time period 1977?2018. Major taxa studied Leaf litter. Methods We synthesized 1,707 records of litter decomposition in streams from 275 studies. We explored variations in decomposition rates among climate zones and tree functional types and between mesh size groups. Regressions were performed to identify the factors that played dominant roles in litter decomposition globally. Results Litter decomposition rates did not differ among tropical, temperate and cold climate zones. Decomposition rates of litter from evergreen conifer trees were much lower than those of deciduous and evergreen broadleaf trees, attributed to the low quality of litter from evergreen conifers. No significant differences were found between decomposition rates of litter from deciduous and evergreen broadleaf trees. Additionally, litter decomposition rates were much higher in coarse- than in fine-mesh bags, which controled the entrance of decomposers of different body sizes. Multiple regressions showed that litter traits (including lignin, C:N ratio) and elevation were the most important factors in regulating leaf litter decomposition. Main conclusions Litter traits predominantly control leaf litter decomposition in streams worldwide. Although further analyses are necessary to explore whether commonalities of the predominant role of litter traits in decomposition exist in both aquatic and terrestrial ecosystems, our findings could contribute to the use of trait-based approaches in modelling the decomposition of litter in streams globally and exploring mechanisms of land?water?atmosphere carbon fluxes.
Zhang Q, Wei W, Chen L, Yang L, Chen HYH, Luo Y (2019) Soil water availability drives changes in community traits along a hydrothermal gradient in Loess Plateau Grasslands. Rangeland Ecology & Management.
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Read PublicationPlant functional traits can be used to predict ecosystem responses to climate gradients, yet precipitation explains very little variation for most traits. Soil water availability directly influences plant water uptake and thus may assist with the improvement of plant trait–water relationships. However, this promise remains poorly realized due to rare tests. Here, we provide the first study that attempts to link climate factors, vertical soil water availability, and community composition at a regional scale. Our study paired field-measured vertical soil available water (0–300 cm) and community functional composition at 46 herbaceous grassland sites along a steep hydrothermal gradient in the Loess Plateau of Central China. Community functional composition was expressed via community-weighted means of eight traits. Structural equation modeling was employed to evaluate the role of vertical soil available water content, controlled by precipitation and air temperature, in affecting plant community-weighted traits. We found that soil available water content at depths of 20–100 cm was typically responsible for mediating the effects of precipitation and air temperature on plant community composition. This emerged as the predominant factor to explain variations in grassland response traits, including leaf area, specific leaf area, and leaf dry matter content. These traits exhibited clear drought-induced shifts along soil desiccation gradients and responded to drier conditions by reducing leaf area/specific leaf area and increasing leaf dry matter content. Our findings rehighlighted soil water availability as the core driver that needs to be considered in the restoration and management of dryland ecosystems.
Zhang X, Manzanedo RD, D'Orangeville L, Rademacher TT, Li J, Bai X, Hou M, Chen Z, Zou F, Song F, Pederson N (2019) Snowmelt and early to mid-growing season water availability augment tree growth during rapid warming in southern Asian boreal forests. Global Change Biolocy.
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Read PublicationBoreal forests are facing profound changes in their growth environment, including warming-induced water deficits, extended growing seasons, accelerated snowmelt, and permafrost thaw. The influence of warming on trees varies regionally, but in most boreal forests studied to date, tree growth has been found to be negatively affected by increasing temperatures. Here, we used a network of Pinus sylvestris tree-ring collections spanning a wide climate gradient the southern end of the boreal forest in Asia to assess their response to climate change for the period 1958?2014. Contrary to findings in other boreal regions, we found that previously negative effects of temperature on tree growth turned positive in the northern portion of the study network after the onset of rapid warming. Trees in the drier portion did not show this reversal in their climatic response during the period of rapid warming. Abundant water availability during the growing season, particularly in the early to mid-growing season (May?July), is key to the reversal of tree sensitivity to climate. Advancement in the onset of growth appears to allow trees to take advantage of snowmelt water, such that tree growth increases with increasing temperatures during the rapidly warming period. The region's monsoonal climate delivers limited precipitation during the early growing season, and thus snowmelt likely covers the water deficit so trees are less stressed from the onset of earlier growth. Our results indicate that the growth response of P. sylvestris to increasing temperatures strongly related to increased early season water availability. Hence, boreal forests with sufficient water available during crucial parts of the growing season might be more able to withstand or even increase growth during periods of rising temperatures. We suspect that other regions of the boreal forest may be affected by similar dynamics.
Zhang X, Manzanedo RD, D'Orangeville L, Rademacher TT, Li J, Bai X, Hou M, Chen Z, Zou F, Song F, Pederson N (2019) Snowmelt and early to mid-growing season water availability augment tree growth during rapid warming in southern Asian boreal forests. Global Change Biology.
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Read PublicationBoreal forests are facing profound changes in their growth environment, including warming-induced water deficits, extended growing seasons, accelerated snowmelt, and permafrost thaw. The influence of warming on trees varies regionally, but in most boreal forests studied to date, tree growth has been found to be negatively affected by increasing temperatures. Here, we used a network of Pinus sylvestris tree-ring collections spanning a wide climate gradient the southern end of the boreal forest in Asia to assess their response to climate change for the period 1958?2014. Contrary to findings in other boreal regions, we found that previously negative effects of temperature on tree growth turned positive in the northern portion of the study network after the onset of rapid warming. Trees in the drier portion did not show this reversal in their climatic response during the period of rapid warming. Abundant water availability during the growing season, particularly in the early to mid-growing season (May?July), is key to the reversal of tree sensitivity to climate. Advancement in the onset of growth appears to allow trees to take advantage of snowmelt water, such that tree growth increases with increasing temperatures during the rapidly warming period. The region's monsoonal climate delivers limited precipitation during the early growing season, and thus snowmelt likely covers the water deficit so trees are less stressed from the onset of earlier growth. Our results indicate that the growth response of P. sylvestris to increasing temperatures strongly related to increased early season water availability. Hence, boreal forests with sufficient water available during crucial parts of the growing season might be more able to withstand or even increase growth during periods of rising temperatures. We suspect that other regions of the boreal forest may be affected by similar dynamics.
Zheng M, Chen H, Li D, Luo Y, Mo J, (2019) Substrate stoichiometry determines nitrogen fixation throughout succession in southern Chinese forests. Ecology Letters.
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Read PublicationThe traditional view holds that biological nitrogen (N) fixation often peaks in early- or mid-successional ecosystems and declines throughout succession based on the hypothesis that soil N richness and/or phosphorus (P) depletion become disadvantageous to N fixers. This view, however, fails to support the observation that N fixers can remain active in many old-growth forests despite the presence of N-rich and/or P-limiting soils. Here, we found unexpected increases in N fixation rates in the soil, forest floor, and moss throughout three successional forests and along six age-gradient forests in southern China. We further found that the variation in N fixation was controlled by substrate carbon(C) : N and C : (N : P) stoichiometry rather than by substrate N or P. Our findings highlight the utility of ecological stoichiometry in illuminating the mechanisms that couple forest succession and N cycling.
Zheng M, Zhang W, Luo Y, Wan S, Fu S, Wang S, Liu N, Ye Q, Yan J, Zou B, Fang C, Ju Y, Ha D, Zhu L,Mo J (2019) The inhibitory effects of nitrogen deposition on asymbiotic nitrogen fixation are divergent between a tropical and a temperate forest. Ecosystems 22(5):955-967.
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Read PublicationAsymbiotic nitrogen (N) fixation (ANF) is an important source of N in pristine forests and is predicted to decrease with N deposition. Previous studies revealing N fixation in response to N deposition have mostly applied understory N addition approaches, neglecting the key processes (for example, N retention and uptake) occurring in forest canopy. This study evaluated the effects of N deposition on N fixation in the soil, forest floor, mosses, and canopy leaves in a temperate forest (in central China) and a tropical forest (in southern China) with different treatments: control, understory N addition, and canopy N addition. Results showed that total ANF rates were higher in the temperate forest (2.57 ± 0.19 mg N m−2 d−1) than in the tropical forest (1.34 ± 0.09 mg N m−2 d−1). N addition inhibited the soil, forest floor, moss, and foliar N fixation in the temperate forest, whereas it inhibited only the soil N fixation in the tropical forest. Compared to canopy N addition, understory N addition overestimated the inhibitory effects of N deposition on total ANF slightly in the tropical forest (by 35%) but severely in the temperate forest (by 375–472%) due to neglecting canopy retention of N. In summary, our findings indicate that ANF has different rates and sensitivities to N addition between tropical and temperate forests and that understory N addition overestimates the N deposition effects on ANF in forests, particularly in the temperate forest. These findings are important for our accurate understanding and estimate of terrestrial N fixation under N deposition scenarios.
Zheng M, Zhou Z, Luo Y, Zhao P, Mo J (2019) Global pattern and controls of biological nitrogen fixation under nutrient enrichment: A meta-analysis. Global Change Biology.
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Read PublicationBiological nitrogen (N) fixation (BNF), an important source of N in terrestrial ecosystems, plays a critical role in terrestrial nutrient cycling and net primary productivity. Currently, large uncertainty exists regarding how nutrient availability regulates terrestrial BNF and the drivers responsible for this process. We conducted a global meta-analysis of terrestrial BNF in response to N, phosphorus (P), and micronutrient (Micro) addition across different biomes (i.e., tropical/subtropical forest, savanna, temperate forest, grassland, boreal forest, and tundra) and explored whether the BNF responses were affected by fertilization regimes (nutrient-addition rates, duration, and total load) and environmental factors (mean annual temperature (MAT), mean annual precipitation (MAP), and N deposition). The results showed that N addition inhibited terrestrial BNF (by 19.0% [95% confidence interval (CI): 17.7?20.3%]; hereafter), Micro addition stimulated terrestrial BNF (30.4% [25.7?35.3%]), and P addition had an inconsistent effect on terrestrial BNF (i.e., inhibiting free-living N fixation (7.5% [4.4?10.6%]) and stimulating symbiotic N fixation (85.5% [25.8?158.7%])). Furthermore, the response ratios (i.e., effect sizes) of BNF to nutrient addition were smaller in low-latitude (<30°) biomes (8.5?36.9%) than in mid-/high-latitude (≥30°) biomes (32.9?61.3%), and the sensitivity (defined as the absolute value of response ratios) of BNF to nutrients in mid-/high-latitude biomes decreased with decreasing latitude (p≤0.009; linear/logarithmic regression models). Fertilization regimes did not affect this phenomenon (p>0.05), but environmental factors did affect it (p<0.001) because MAT, MAP, and N deposition accounted for 5?14%, 10?32%, and 7?18% of the variance in the BNF response ratios in cold (MAT<15°C), low-rainfall (MAP<2500 mm), and low-N-deposition (<7 kg ha?1 yr?1) biomes, respectively. Overall, our meta-analysis depicts a global pattern of nutrient impacts on terrestrial BNF and indicates that certain types of global change (i.e., warming, elevated precipitation and N deposition) may reduce the sensitivity of BNF in response to nutrient enrichment in mid-/high-latitude biomes. This article is protected by copyright.
Zhou W, Guo Z, Chen J, Jiang J, Hui D, Wang X, Sheng J, Chen L, Luo Y, Zheng J, Li S, Zhang Y (2019) Direct seeding for rice production increased soil erosion and phosphorus runoff losses in subtropical China. Science of the Total Environment 695: 133845.
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Read PublicationEstimating soil erosion and nutrient losses from surface runoff in paddy fields is essential for the assessment of sustainable rice (Oryza sativa L.) production and water quality protection. Different rice establishment methods have been used in the last three decades in Asia; however, it is still unclear how these methods influence sustainable agriculture and environmental protection in humid areas. The aim of this study was to evaluate the impacts of rice establishment method on soil erosion and phosphorus (P) losses from surface runoff in Hydragric Anthrosols under a subtropical monsoon climate. Total suspended solids (TSS), total P (TP), dissolved P (DP), and particulate P (PP) runoff losses were measured under four rice establishment treatments in 2013 and 2014, including traditional manual transplanting (TT), mechanical transplanting (MT), dry direct seeding (DD), and wet direct seeding (WD). The results showed that the seasonal TSS in the runoff varied from 59.9 to 829.8 kg ha−1 in the two years. Compared with TT, the DD significantly increased the TSS by 481% in 2013 and by 349% in 2014, while the WD significantly increased TSS by 783% in 2013 and by 571% in 2014. In the 2013 and 2014 rice seasons, the field-observed TP runoff losses were from 0.18 to 1.51 kg ha−1. Compared with TT, the DD significantly increased the TP lost by 222% in 2013 and by 197% in 2014, whereas the WD significantly increased the TP lost by 483% in 2013 and by 387% in 2014. However, the TSS and P losses from the MT and TT were similar in both years. The PP runoff losses accounted for 58–77% of the seasonal TP lost. These findings demonstrate that the conversion of traditional manual transplanting to direct seeding increased soil erosion and P runoff losses in subtropical China.
2018
Adair EC, Hooper DU, Paquette A, Hungate BA (2018) Ecosystem context illuminates conflicting roles of plant diversity in carbon storage. Ecology Letters 21(11): 1604-1619.
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Read PublicationPlant diversity can increase biomass production in plot-scale studies, but applying these results to ecosystem carbon (C) storage at larger spatial and temporal scales remains problematic. Other ecosystem controls interact with diversity and plant production, and may influence soil pools differently from plant pools. We integrated diversity with the state-factor framework, which identifies key controls, or ?state factors?, over ecosystem properties and services such as C storage. We used this framework to assess the effects of diversity, plant traits and state factors (climate, topography, time) on live tree, standing dead, organic horizon and total C in Québec forests. Four patterns emerged: (1) while state factors were usually the most important model predictors, models with both state and biotic factors (mean plant traits and diversity) better predicted C pools; (2) mean plant traits were better predictors than diversity; (3) diversity increased live tree C but reduced organic horizon C; (4) different C pools responded to different traits and diversity metrics. These results suggest that, where ecosystem properties result from multiple processes, no simple relationship may exist with any one organismal factor. Integrating biodiversity into ecosystem ecology and assessing both traits and diversity improves our mechanistic understanding of biotic effects on ecosystems.
Alexander HD, Natali SM, Loranty MM, Ludwig SM, Spektor VV, Davydov S, Zimov N, Trujillo I, Mack MC (2018) Impacts of increased soil burn severity on larch forest regeneration on permafrost soils of far northeastern Siberia. Forest Ecology and Management 417: 144-153.
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Read PublicationFire severity is increasing across the boreal forest biome as climate warms, and initial post-fire changes in tree demographic processes could be important determinants of long-term forest structure and carbon dynamics. To examine soil burn severity impacts on tree regeneration, we conducted experimental burns in summer 2012 that created a gradient of residual post-fire soil organic layer (SOL) depth within a mature, sparse-canopy Cajander larch (Larix cajanderi Mayr.) forest in the Eastern Siberian Arctic. Each fall from 2012 to 2016, we added larch seeds to plots along the burn severity gradient. We tracked density of new larch germinants and established seedlings (alive ≥ 1 year) during subsequent growing seasons, along with changes in seedbed conditions (permafrost thaw depth, moisture, and temperature). Over the study, a cumulative total of 17 and 18 new germinants m−2 occurred in high and moderate severity treatments, respectively, while germinants were rare in unburned and low severity treatments (< 0.5 germinants m−2). Most seedlings (> 50%) germinated in summer 2017, following a mast event in fall 2016, suggesting safe sites for germination were not fully occupied in previous years despite seed additions. By 2017, established seedling density was ∼5 times higher on moderate and high severity treatments compared to other treatments. Cumulative total density of new germinants and established seedlings increased linearly with decreasing residual SOL depth, as did thaw depth, soil moisture, and soil temperature. Our findings suggest that increased soil burn severity could improve seedbed conditions and increase larch recruitment, assuming seed sources are available. If these demographic changes persist as stands mature, a climate-driven increase in soil burn severity could shift forest structure from sparse-canopy stands, which dominate this region of the Siberian Arctic, to high density stands, with potential implications for carbon, energy, and water cycling.
Bjorkman AD, Myers-Smith IH, Elmendorf SC, Normand S, Thomas HJD, Alatalo JM, Alexander H, Anadon-Rosell A, Angers-Blondin S, Bai Y, Baruah G, te Beest M, Berner L, Bjork RG, Blok D, Bruelheide H, Buchwal A, Buras A, Carbognani M, Christie K, Collier LS, Cooper EJ, Cornelissen JHC, Dickinson KJM, Dullinger S, Elberling B, Eskelinen A, Forbes BC, Frei ER, Iturrate-Garcia M, Good MK, Grau O, Green P, Greve M, Grogan P, Haider S, Hajek T, Hallinger M, Happonen K, Harper KA, Heijmans MMPD, Henry GHR, Hermanutz L, Hewitt RE, Hollister RD, Hudson J, Hulber K, Iversen CM, Jaroszynska F, Jimenez-Alfaro B, Johnstone J, Jorgensen RH, Kaarlejarvi E, Klady R, Klimesova J, Korsten A, Kuleza S, Kulonen A, Lamarque LJ, Lantz T, Lavalle A, Lebrechts JJ, Levesque E, Little CJ, Luoto M, Macek P, Mack MC, Mathakutha R, MIchelsen A, Milbau A, Molau U, Morgan JW, Morsdorf MA, Nabe-Nielsen J, Nielsen SS, Ninot JM, Oberbauer SF, Olofsson J, Onipchenko VG, Petraglia A, Pickering C, Pevey JS, Rixen C, Rumpf SB, Schaepman-Strub G, Semenchuk P, Shetti R, Soudzilovskaia NA, Spasojevic MJ, Speed JDM, Street LE, Suding K, Tape KD, Tomaselli M, Trant A, Treier UA, Tremblay J, Tremblay M, Venn S, Virkkala AM, Vowles T, Weijers S, Wilmking M, Wipf S, Zamin T (2018) Tundra Trait Team: A database of plant traits spanning the tundra biome. Global Ecology and Biogeography 27(12): 1402-1411.
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Read PublicationThe Tundra Trait Team (TTT) database includes field-based measurements of key traits related to plant form and function at multiple sites across the tundra biome. This dataset can be used to address theoretical questions about plant strategy and trade-offs, trait?environment relationships and environmental filtering, and trait variation across spatial scales, to validate satellite data, and to inform Earth system model parameters. Main types of variable contained The database contains 91,970 measurements of 18 plant traits. The most frequently measured traits (>1,000 observations each) include plant height, leaf area, specific leaf area, leaf fresh and dry mass, leaf dry matter content, leaf nitrogen, carbon and phosphorus content, leaf C:N and N:P, seed mass, and stem specific density. Spatial location and grain Measurements were collected in tundra habitats in both the Northern and Southern Hemispheres, including Arctic sites in Alaska, Canada, Greenland, Fennoscandia and Siberia, alpine sites in the European Alps, Colorado Rockies, Caucasus, Ural Mountains, Pyrenees, Australian Alps, and Central Otago Mountains (New Zealand), and sub-Antarctic Marion Island. More than 99% of observations are georeferenced. Time period and grain All data were collected between 1964 and 2018. A small number of sites have repeated trait measurements at two or more time periods. Major taxa and level of measurement Trait measurements were made on 978 terrestrial vascular plant species growing in tundra habitats. Most observations are on individuals (86%), while the remainder represent plot or site means or maximums per species. Software format csv file and GitHub repository with data cleaning scripts in R; contribution to TRY plant trait database (www.try-db.org) to be included in the next version release.
Blankinship JC, Berhe AA, Crow SE, Druhan JL, Heckman KA, Keiluweit M, Lawrence CR, Marin-Spiotta E, Plante AF, Rasmussen C, Schädel C, Schimel JP, Sierra CA, Thompson A, Wagai R, Weider WR (2018) Improving understanding of soil organic matter dynamics by triangulating theories, measurements, and models. Biogeochemistry 140(1): 1-13.
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Read PublicationSoil organic matter (SOM) turnover increasingly is conceptualized as a tension between accessibility to microorganisms and protection from decomposition via physical and chemical association with minerals in emerging soil biogeochemical theory. Yet, these components are missing from the original mathematical models of belowground carbon dynamics and remain underrepresented in more recent compartmental models that separate SOM into discrete pools with differing turnover times. Thus, a gap currently exists between the emergent understanding of SOM dynamics and our ability to improve terrestrial biogeochemical projections that rely on the existing models. In this opinion paper, we portray the SOM paradigm as a triangle composed of three nodes: conceptual theory, analytical measurement, and numerical models. In successful approaches, we contend that the nodes are connected—models capture the essential features of dominant theories while measurement tools generate data adequate to parameterize and evaluate the models—and balanced—models can inspire new theories via emergent behaviors, pushing empiricists to devise new measurements. Many exciting advances recently pushed the boundaries on one or more nodes. However, newly integrated triangles have yet to coalesce. We conclude that our ability to incorporate mechanisms of microbial decomposition and physicochemical protection into predictions of SOM change is limited by current disconnections and imbalances among theory, measurement, and modeling. Opportunities to reintegrate the three components of the SOM paradigm exist by carefully considering their linkages and feedbacks at specific scales of observation.
Brown CD, Dufour-Tremblay G, Jameson RG, Mamet SD, Trant AJ, Walker XJ, Boudreau S, Harper KA, Henry GHR, Hermanutz L, Hofgaard A, Isaeva L, Kershaw GP, Johnstone JF (2018) Reproduction as a bottleneck to treeline advance across the circumarctic forest tundra ecotone. Ecography.
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Read PublicationThe fundamental niche of many species is shifting with climate change, especially in sub-arctic ecosystems with pronounced recent warming. Ongoing warming in sub-arctic regions should lessen environmental constraints on tree growth and reproduction, leading to increased success of trees colonising tundra. Nevertheless, variable responses of treeline ecotones have been documented in association with warming temperatures. One explanation for time lags between increasingly favourable environmental conditions and treeline ecotone movement is reproductive limitations caused by low seed availability. Our objective was to assess the reproductive constraints of the dominant tree species at the treeline ecotone in the circumpolar north. We sampled reproductive structures of trees (cones and catkins) and stand attributes across circumarctic treeline ecotones. We used generalized linear mixed models to estimate the sensitivity of seed production and the availability of viable seed to regional climate, stand structure, and species-specific characteristics. Both seed production and viability of available seed were strongly driven by specific, sequential seasonal climatic conditions, but in different ways. Seed production was greatest when growing seasons with more growing degree days coincided with years with high precipitation. Two consecutive years with more growing degree days and low precipitation resulted in low seed production. Seasonal climate effects on the viability of available seed depended on the physical characteristics of the reproductive structures. Large-coned and -seeded species take more time to develop mature embryos and were therefore more sensitive to increases in growing degree days in the year of flowering and embryo development. Our findings suggest that both moisture stress and abbreviated growing seasons can have a notable negative influence on the production and viability of available seed at treeline. Our synthesis revealed that constraints on predispersal reproduction within the treeline ecotone might create a considerable time lag for range expansion of tree populations into tundra ecosystems.
Buermann W, Forkel M, O'Sullivan M, Sitch S, Friedlingstein P, Haverd V, Jain AK, Kato E, Kautz M, Lienert S, Lombardozzi D, Nabel JEMS, Tian H, Wiltshire AJ, Zhu D, Smith WK, Richardson AD (2018) Widespread seasonal compensation effects of spring warming on northern plant productivity. Nature 562(7725): 110-114.
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Read PublicationClimate change is shifting the phenological cycles of plants1, thereby altering the functioning of ecosystems, which in turn induces feedbacks to the climate system2. In northern (north of 30° N) ecosystems, warmer springs lead generally to an earlier onset of the growing season3,4 and increased ecosystem productivity early in the season5. In situ6 and regional7–9 studies also provide evidence for lagged effects of spring warmth on plant productivity during the subsequent summer and autumn. However, our current understanding of these lagged effects, including their direction (beneficial or adverse) and geographic distribution, is still very limited. Here we analyse satellite, field-based and modelled data for the period 1982–2011 and show that there are widespread and contrasting lagged productivity responses to spring warmth across northern ecosystems. On the basis of the observational data, we find that roughly 15 per cent of the total study area of about 41 million square kilometres exhibits adverse lagged effects and that roughly 5 per cent of the total study area exhibits beneficial lagged effects. By contrast, current-generation terrestrial carbon-cycle models predict much lower areal fractions of adverse lagged effects (ranging from 1 to 14 per cent) and much higher areal fractions of beneficial lagged effects (ranging from 9 to 54 per cent). We find that elevation and seasonal precipitation patterns largely dictate the geographic pattern and direction of the lagged effects. Inadequate consideration in current models of the effects of the seasonal build-up of water stress on seasonal vegetation growth may therefore be able to explain the differences that we found between our observation-constrained estimates and the model-constrained estimates of lagged effects associated with spring warming. Overall, our results suggest that for many northern ecosystems the benefits of warmer springs on growing-season ecosystem productivity are effectively compensated for by the accumulation of seasonal water deficits, despite the fact that northern ecosystems are thought to be largely temperature- and radiation-limited10.
Cai A, Liang G, Zhang X, Zhang W, Li L Rui Y, Xu M, Luo Y (2018) Long-term straw decomposition in agro-ecosystems described by a unified three-exponentiation equation with thermal time. Science of the total environment 636: 699-708.
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Read PublicationUnderstanding drivers of straw decomposition is essential for adopting appropriate management practice to improve soil fertility and promote carbon (C) sequestration in agricultural systems. However, predicting straw decomposition and characteristics is difficult because of the interactions between many factors related to straw properties, soil properties, and climate, especially under future climate change conditions. This study investigated the driving factors of straw decomposition of six types of crop straw including wheat, maize, rice, soybean, rape, and other straw by synthesizing 1642 paired data from 98 published papers at spatial and temporal scales across China. All the data derived from the field experiments using little bags over twelve years. Overall, despite large differences in climatic and soil properties, the remaining straw carbon (C, %) could be accurately represented by a three-exponent equation with thermal time (accumulative temperature). The lignin/nitrogen and lignin/phosphorus ratios of straw can be used to define the size of labile, intermediate, and recalcitrant C pool. The remaining C for an individual type of straw in the mild-temperature zone was higher than that in the warm-temperature and subtropical zone within one calendar year. The remaining straw C after one thermal year was 40.28%, 37.97%, 37.77%, 34.71%, 30.87%, and 27.99% for rice, soybean, rape, wheat, maize, and other straw, respectively. Soil available nitrogen and phosphorus influenced the remaining straw C at different decomposition stages. For one calendar year, the total amount of remaining straw C was estimated to be 29.41 Tg and future temperature increase of 2 °C could reduce the remaining straw C by 1.78 Tg. These findings confirmed the long-term straw decomposition could be mainly driven by temperature and straw quality, and quantitatively predicted by thermal time with the three-exponent equation for a wide array of straw types at spatial and temporal scales in agro-ecosystems of China.
Ceballos-Nunez V, Richardson AD, Sierra CA (2018) Ages and transit times as important diagnostics of model performance for predicting carbon dynamics in terrestrial vegetation models. Biogeosciences 15: 1607-1625.
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Read PublicationThe global carbon cycle is strongly controlled by the source/sink strength of vegetation as well as the capacity of terrestrial ecosystems to retain this carbon. These dynamics, as well as processes such as the mixing of old and newly fixed carbon, have been studied using ecosystem models, but different assumptions regarding the carbon allocation strategies and other model structures may result in highly divergent model predictions. We assessed the influence of three different carbon allocation schemes on the C cycling in vegetation. First, we described each model with a set of ordinary differential equations. Second, we used published measurements of ecosystem C compartments from the Harvard Forest Environmental Measurement Site to find suitable parameters for the different model structures. And third, we calculated C stocks, release fluxes, radiocarbon values (based on the bomb spike), ages, and transit times. We obtained model simulations in accordance with the available data, but the time series of C in foliage and wood need to be complemented with other ecosystem compartments in order to reduce the high parameter collinearity that we observed, and reduce model equifinality. Although the simulated C stocks in ecosystem compartments were similar, the different model structures resulted in very different predictions of age and transit time distributions. In particular, the inclusion of two storage compartments resulted in the prediction of a system mean age that was 12–20 years older than in the models with one or no storage compartments. The age of carbon in the wood compartment of this model was also distributed towards older ages, whereas fast cycling compartments had an age distribution that did not exceed 5 years. As expected, models with C distributed towards older ages also had longer transit times. These results suggest that ages and transit times, which can be indirectly measured using isotope tracers, serve as important diagnostics of model structure and could largely help to reduce uncertainties in model predictions. Furthermore, by considering age and transit times of C in vegetation compartments as distributions, not only their mean values, we obtain additional insights into the temporal dynamics of carbon use, storage, and allocation to plant parts, which not only depends on the rate at which this C is transferred in and out of the compartments but also on the stochastic nature of the process itself.
Chen J, Luo Y, Garcia-Palacios P, Cao J, Dacal M, Zhou X, Li J, Xia J, Niu S, Yang H, Shelton S, Guo W, van Groenigen KJ (2018) Differential responses of carbon-degrading enzyme activities to warming: Implications for soil respiration. Global Change Biology 24(10):4816-4826.
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Read PublicationExtracellular enzymes catalyze rate-limiting steps in soil organic matter decomposition, and their activities (EEAs) play a key role in determining soil respiration (SR). Both EEAs and SR are highly sensitive to temperature, but their responses to climate warming remain poorly understood. Here, we present a meta-analysis on the response of soil cellulase and ligninase activities and SR to warming, synthesizing data from 56 studies. We found that warming significantly enhanced ligninase activity by 21.4% but had no effect on cellulase activity. Increases in ligninase activity were positively correlated with changes in SR, while no such relationship was found for cellulase. The warming response of ligninase activity was more closely related to the responses of SR than a wide range of environmental and experimental methodological factors. Furthermore, warming effects on ligninase activity increased with experiment duration. These results suggest that soil microorganisms sustain long-term increases in SR with warming by gradually increasing the degradation of the recalcitrant carbon pool.
Chen J, Luo Y, van Groenigen KJ, Hungate BA, Cao J, Zhou X, Wang R (2018) A keystone microbial enzyme for nitrogen control of soil carbon storage. Science Advances 4(8): eaaq1689.
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Read PublicationAgricultural and industrial activities have increased atmospheric nitrogen (N) deposition to ecosystems worldwide. N deposition can stimulate plant growth and soil carbon (C) input, enhancing soil C storage. Changes in microbial decomposition could also influence soil C storage, yet this influence has been difficult to discern, partly because of the variable effects of added N on the microbial enzymes involved. We show, using meta-analysis, that added N reduced the activity of lignin-modifying enzymes (LMEs), and that this N-induced enzyme suppression was associated with increases in soil C. In contrast, N-induced changes in cellulase activity were unrelated to changes in soil C. Moreover, the effects of added soil N on LME activity accounted for more of the variation in responses of soil C than a wide range of other environmental and experimental factors. Our results suggest that, through responses of a single enzyme system to added N, soil microorganisms drive long-term changes in soil C accumulation. Incorporating this microbial influence on ecosystem biogeochemistry into Earth system models could improve predictions of ecosystem C dynamics.
Chen J, Luo Y, Xia J, Zhou X, Niu S, Shelton S, Guo W, Liu S, Dai W, Cao J (2018) Divergent responses of ecosystem respiration components to livestock exclusion on the Quighai Tibetan Plateau. Land Degradation & Development 29(6):1726-1737.
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Read PublicationGrazing exclusion (GE) is an effective method for protecting degraded grasslands, and it can profoundly affect ecosystem carbon (C) cycles. Ecosystem respiration (ER), which includes both autotrophic and heterotrophic respiration (HR), accounts for the largest land-to-atmosphere C fluxes. How ER responds to GE is still unclear, however, and to investigate this, a controlled GE experiment was conducted at a meadow grassland near Qinghai Lake, China. Animal exclusion enhanced ER and aboveground plant respiration (Ragb) by 10.5% and 40.1%, respectively, but it suppressed soil respiration by 12.4% and HR by 17.6%. Positive responses of ER and Ragb were linked to increased aboveground biomass, particularly graminoids biomass. Negative responses of soil respiration and HR were associated with GE-induced changes in microbial biomass C and nitrogen. These results show that grassland responded in complex ways to GE and that ER and its components were regulated by both abiotic and biotic factors. Moreover, the divergent responses of respiration components have important implications for models of terrestrial C cycles and climate under enhanced human activities and changes in land use.
Chen L, Sun L, Liu W, Wang L, Wu H, Zhu AX, Luo Y (2018) Evapotranspiration partitioning using an optimality-based ecohydrological model in a semiarid shrubland. International Journal of Digital Earth.
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Read PublicationPartitioning of evapotranspiration (ET) into biological component transpiration (T) and non-biological component evaporation (E) is crucial in understanding the impact of environmental change on ecosystems and water resources. However, direct measurement of transpiration is still challenging. In this paper, an optimality-based ecohydrological model named Vegetation Optimality Model (VOM) is applied for ET partitioning. The results show that VOM model can reasonably simulate ET and ET components in a semiarid shrubland. Overall, the ratio of transpiration to evapotranspiration is 49% for the whole period. Evaporation and plant transpiration mainly occur in monsoon following the precipitation events. Evaporation responds immediately to precipitation events, while transpiration shows a lagged response of several days to those events. Different years demonstrate different patterns of T/ET ratio dynamic in monsoon. Some of the years show a low T/ET ratio at the beginning of monsoon and slowly increased T/ET ratio. Other years show a high level of T/ET ratio for the whole monsoon. We find out that spring precipitation, especially the size of the precipitation, has a significant influence on the T/ET ratio in monsoon.
Christiansen CT, Mack MC, DeMarco J, Grogan P (2018) Decomposition of Senesced leaf litter is faster in tall compared to low birch shrub tundra. Ecosystems 1-16.
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Read PublicationMany Low Arctic tundra regions are currently undergoing a vegetation shift towards increasing growth and groundcover of tall deciduous shrubs due to recent climate warming. Vegetation change directly affects ecosystem carbon balance, but it can also affect soil biogeochemical cycling through physical and biological feedback mechanisms. Recent studies indicate that enhanced snow accumulation around relatively tall shrubs has negligible physical effect on litter decomposition rates. However, these investigations were no more than 3 years, and therefore may be insufficient to detect differences in inherently slow biogeochemical processes. Here, we report a 5-year study near Daring Lake, Canada, comparing Betula neoalaskana foliar litter decay rates within unmanipulated and snowfenced low-stature birch (height: ~ 0.3 m) plots to test the physical effect of experimentally deepened snow, and within tall birch (height: ~ 0.8 m) plots to test the combined physical and biological effects, that is, deepened snow plus strong birch dominance. Having corrected for carbon gain by the colonizing decomposers, actual litter carbon loss increased by approximately 25% in the tall birch relative to both low birch sites. Decay of lignin-like acid unhydrolizable litter residues also accelerated in the tall birch site, and a similar but lower magnitude response in the snowfenced low birch site indicated that physical effects of deepened snow were at least partially responsible. In contrast, deepened snow alone did not affect litter carbon loss. Our findings suggest that a combination of greater litter inputs, altered soil microbial community, enhanced soil nutrient pools, and warmer winter soils together promote relatively fast decomposition of recalcitrant litter carbon in tall birch shrub environments.
Compson ZG, Hungate BA, Whitham TG, Koch GW, Dijkstra P, Siders AC, Wojtowicz T, Jacobs R, Rakestraw DN, Allred KE, Sayer Ck, Marks JC (2018) Linking tree genetics and stream consumers: Isotopic tracers elucidate controls on carbon and nitrogen assimilation. Ecology.
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Read PublicationLeaf litter provides an important nutrient subsidy to headwater streams, but little is known about how tree genetics influences energy pathways from litter to higher trophic levels. Despite the charge to quantify carbon (C) and nitrogen (N) pathways from decomposing litter, the relationship between litter decomposition and aquatic consumers remains unresolved. We measured litter preference (attachments to litter), C and N assimilation rates, and growth rates of a shredding caddisfly (Hesperophylax magnus, Limnephilidae) in response to leaf litter of different chemical and physical phenotypes using Populus cross types (P. fremontii, P. angustifolia, and F1 hybrids) and genotypes within P. angustifolia. We combined laboratory mesocosm studies using litter from a common garden with a field study using doubly labeled litter (13 C and 15 N) grown in a greenhouse and incubated in Oak Creek, AZ. We found that, in the lab, shredders initially chose relatively labile (low lignin and condensed tannin concentrations, rapidly decomposing) cross type litter, but preference changed within four days to relatively recalcitrant (high lignin and condensed tannin concentrations, slowly decomposing) litter types. Additionally, in the lab, shredder growth rates were higher on relatively recalcitrant compared to labile cross type litter. Over the course of a three-week field experiment, shredders also assimilated more C and N from relatively recalcitrant compared to labile cross type litter. Finally, among P. angustifolia genotypes, N assimilation by shredders was positively related to litter lignin and C:N, but negatively related to condensed tannins and decomposition rate. C assimilation was likewise positively related to litter C:N, and also to litter %N. C assimilation was not associated with condensed tannins or lignin. Collectively, these findings suggest that relatively recalcitrant litter of Populus cross types provides more nutritional benefit, in terms of N fluxes and growth, than labile litter, but among P. angustifolia genotypes the specific trait of litter recalcitrance (lignin or tannins) determines effects on C or N assimilation. As shredders provide nutrients and energy to higher trophic levels, the influence of these genetically based plant decomposition pathways on shredder preference and performance may affect community and food web structure.
Davis GS, Waits K, Nordstrom L, Grande H, Weaver B, Papp K, Horwinski J, Koch B, Hungate BA, Liu CM, Price LB (2018) Antibiotic-resistant Escherichia coli from retail poultry meat with different antibiotic use claims. BMC Microbiology 18(1): 174.
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Read PublicationWe sought to determine if the prevalence of antibiotic-resistant Escherichia coli differed across retail poultry products and among major production categories, including organic, “raised without antibiotics”, and conventional.
Du L, Mikle N, Zou Z, Huang Y, Shi Z, Jiang L, McCarthy HR, Liang J, Luo Y (2018) Global patterns of extreme drought-induced loss in land primary production: Identifying ecological extremes from rain-use efficiency. The Science of the total environment 628-629/611-620.
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Read PublicationQuantifying the ecological patterns of loss of ecosystem function in extreme drought is important to understand the carbon exchange between the land and atmosphere. Rain-use efficiency [RUE; gross primary production (GPP)/precipitation] acts as a typical indicator of ecosystem function. In this study, a novel method based on maximum rain-use efficiency (RUEmax) was developed to detect losses of ecosystem function globally. Three global GPP datasets from the MODIS remote sensing data (MOD17), ground upscaling FLUXNET observations (MPI-BGC), and process-based model simulations (BESS), and a global gridded precipitation product (CRU) were used to develop annual global RUE datasets for 2001-2011. Large, well-known extreme drought events were detected, e.g. 2003 drought in Europe, 2002 and 2011 drought in the U.S., and 2010 drought in Russia. Our results show that extreme drought-induced loss of ecosystem function could impact 0.9% ± 0.1% of earth's vegetated land per year and was mainly distributed in semi-arid regions. The reduced carbon uptake caused by functional loss (0.14 ± 0.03 PgC/yr) could explain >70% of the interannual variation in GPP in drought-affected areas (p ≤ 0.001). Our results highlight the impact of ecosystem function loss in semi-arid regions with increasing precipitation variability and dry land expansion expected in the future. Copyright © 2018 Elsevier B.V. All rights reserved.
Du Z, Weng E, Jiang L, Luo Y, Jianyang X, Zhou X (2018) Carbon–nitrogen coupling under three schemes of model representation: a traceability analysis. Geoscientific Model Development 11: 4399-4416.
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Read PublicationThe interaction between terrestrial carbon (C) and nitrogen (N) cycles has been incorporated into more and more land surface models. However, the scheme of C–N coupling differs greatly among models, and how these diverse representations of C–N interactions will affect C-cycle modeling remains unclear. In this study, we explored how the simulated ecosystem C storage capacity in the terrestrial ecosystem (TECO) model varied with three different commonly used schemes of C–N coupling. The three schemes (SM1, SM2, and SM3) have been used in three different coupled C–N models (i.e., TECO-CN, CLM 4.5, and O-CN, respectively). They differ mainly in the stoichiometry of C and N in vegetation and soils, plant N uptake strategies, downregulation of photosynthesis, and the pathways of N import. We incorporated the three C–N coupling schemes into the C-only version of the TECO model and evaluated their impacts on the C cycle with a traceability framework. Our results showed that all three of the C–N schemes caused significant reductions in steady-state C storage capacity compared with the C-only version with magnitudes of <span class="inline-formula">−23</span> %, <span class="inline-formula">−30</span> %, and <span class="inline-formula">−54</span> % for SM1, SM2, and SM3, respectively. This reduced C storage capacity was mainly derived from the combined effects of decreases in net primary productivity (NPP; <span class="inline-formula">−29</span> %, <span class="inline-formula">−15</span> %, and <span class="inline-formula">−</span>45 %) and changes in mean C residence time (MRT; 9 %, <span class="inline-formula">−17</span> %, and <span class="inline-formula">−17</span> %) for SM1, SM2, and SM3, respectively. The differences in NPP are mainly attributed to the different assumptions on plant N uptake, plant tissue <span class="inline-formula">C</span> <span class="inline-formula">:</span> <span class="inline-formula">N</span> ratio, downregulation of photosynthesis, and biological N fixation. In comparison, the alternative representations of the plant vs. microbe competition strategy and the plant N uptake, combined with the flexible <span class="inline-formula">C</span> <span class="inline-formula">:</span> <span class="inline-formula">N</span> ratio in vegetation and soils, led to a notable spread in MRT. These results highlight the fact that the diverse assumptions on N processes represented by different C–N coupled models could cause additional uncertainty for land surface models. Understanding their difference can help us improve the capability of models to predict future biogeochemical cycles of terrestrial ecosystems.
Espinosa S, Celis G, Branch LC (2018) When roads appear jaguars decline: Increased access to an Amazonian wilderness area reduces potential for jaguar conservation. PLOS ONE 12(1): e0189740.
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Read PublicationRoads are a main threat to biodiversity conservation in the Amazon, in part, because roads increase access for hunters. We examine how increased landscape access by hunters may lead to cascading effects that influence the prey community and abundance of the jaguar (Panthera onca), the top Amazonian terrestrial predator. Understanding such ecological effects originating from anthropogenic actions is essential for conservation and management of wildlife populations in areas undergoing infrastructure development. Our study was conducted in Yasuní Biosphere Reserve, the protected area with highest potential for jaguar conservation in Ecuador, and an area both threatened by road development and inhabited by indigenous groups dependent upon bushmeat. We surveyed prey and jaguar abundance with camera traps in four sites that differed in accessibility to hunters and used site occupancy and spatially explicit capture-recapture analyses to evaluate prey occurrence and estimate jaguar density, respectively. Higher landscape accessibility to hunters was linked with lower occurrence and biomass of game, particularly white-lipped peccary (Tayassu pecari) and collared peccary (Pecari tajacu), the primary game for hunters and prey for jaguars. Jaguar density was up to 18 times higher in the most remote site compared to the most accessible site. Our results provide a strong case for the need to: 1) consider conservation of large carnivores and other wildlife in policies about road construction in protected areas, 2) coordinate conservation initiatives with local governments so that development activities do not conflict with conservation objectives, and 3) promote development of community-based strategies for wildlife management that account for the needs of large carnivores.
Fell M, Barber J, Lichstein JW, Ogle K (2018) Multidimensional trait space informed by a mechanistic model of tree growth and carbon allocation. Ecosphere 9(1): e02060.
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Read PublicationPlant functional traits research has revealed many interesting and important patterns among morphological, physiological, and life-history traits and the environment. These are exemplified in trade-offs between groups of traits such as those embodied in the leaf and wood economics spectra. Inferences from empirical studies are often constrained by the correlative nature of the analyses, availability of trait data, and a focus on easily measured traits. However, empirical studies have been fundamental to modeling endeavors aiming to enhance our understanding of how functional traits scale up to affect, for example, community dynamics and ecosystem productivity. Here, we take a complementary approach utilizing an individual-based model of tree growth and mortality (the allometrically constrained growth and carbon allocation [ACGCA] model) to investigate the theoretical trait space (TTS) of North American trees. The model includes 32 parameters representing allometric, physiological, and anatomical traits, some overlapping leaf and wood economics spectra traits. Using a Bayesian approach, we fit the ACGCA model to individual tree heights and diameters from the USFS Forest Inventory and Analysis (FIA) dataset, with further constraints by literature-based priors. Fitting the model to 1.3 million FIA records?aggregated across individuals, species, and sites?produced a posterior distribution of traits leading to realistic growth. We explored this multidimensional posterior distribution (the TTS) to evaluate trait?trait relationships emerging from the ACGCA model, and compare these against empirical patterns reported in the literature. Only three notable bivariate correlations, among 496 possible trait pairs, were contained in the TTS. However, stepwise regressions uncovered a complicated structure; only a subset of traits?related to photosynthesis (e.g., radiation-use efficiency and maintenance respiration)?exhibited strong multivariate trade-offs with each other, while half of the traits?mostly related to allometries and construction costs?varied independently of other traits. Interestingly, specific leaf area was related to several rarely measured root traits. The trade-offs contained in the TTS generally reflect mass-balance (related to carbon allocation) and engineering (mostly related to allometries) trade-offs represented in the ACGCA model and point to potentially important traits that are under-explored in field studies (e.g., root traits and branch senescence rates).
Fell M, Ogle K (2018) Refinement of a theoretical trait space for North American trees via environmental filtering. Ecological Monographs 88(3): 372-384.
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Read PublicationWe refer to a theoretical trait space (TTS) as an n-dimensional hypervolume (hypercube) characterizing the range of values and covariations among multiple functional traits, in the absence of explicit filtering mechanisms. We previously constructed a 32-dimensional TTS for North American trees by fitting the Allometrically Constrained Growth and Carbon Allocation (ACGCA) model to USFS Forest Inventory and Analysis (FIA) data. Here, we sampled traits from this TTS, representing different individual ?trees,? and subjected these trees to a series of gap dynamics simulations resulting in different annual light levels to explore the impact of environmental filtering (light stress) on the trait space. Variation in light limitation led to non-random mortality and a refinement of the TTS. We investigated potential mechanisms underlying such filtering processes by exploring how traits and the environment relate to mortality rates at the tree, phenotype (a specific set of trait values), and stand (a specific gap scenario) levels. The average light level at the forest floor explained 42% of the stand-level mortality, while phenotype- and tree-level mortality were best explained by six functional traits, especially radiation-use efficiency, maximum tree height, and xylem conducting area to sapwood area ratio (?X). These six ?mortality? traits and six traits related to the leaf and wood economics spectra were used to construct trait hypercubes represented by trees that died or survived each gap scenario. For trees that survived, the volume of their refined trait space decreased linearly with increasing stand-level mortality (up to ~50% mortality); the location also shifted, as indicated by non-zero distances between the hypercube centroids of surviving trees compared to dead trees and the original TTS. Overall, the patterns were consistent with empirical studies of functional traits, in terms of which traits predict mortality and the direction of the relationships. This work, however, also identified potentially important functional traits that are not commonly measured in empirical studies, such as ?X and senescence rates of relatively long-lived tissues.
Fer I, Kelly R, Moorcroft PR, Richardson AD, Cowdery EM, Dietze MC (2018) Linking big models to big data: efficient ecosystem model calibration through Bayesian model emulation. Biogeosciences 15: 5801-5830.
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Read PublicationData-model integration plays a critical role in assessing and improving our capacity to predict ecosystem dynamics. Similarly, the ability to attach quantitative statements of uncertainty around model forecasts is crucial for model assessment and interpretation and for setting field research priorities. Bayesian methods provide a rigorous data assimilation framework for these applications, especially for problems with multiple data constraints. However, the Markov chain Monte Carlo (MCMC) techniques underlying most Bayesian calibration can be prohibitive for computationally demanding models and large datasets. We employ an alternative method, Bayesian model emulation of sufficient statistics, that can approximate the full joint posterior density, is more amenable to parallelization, and provides an estimate of parameter sensitivity. Analysis involved informative priors constructed from a meta-analysis of the primary literature and specification of both model and data uncertainties, and it introduced novel approaches to autocorrelation corrections on multiple data streams and emulating the sufficient statistics surface. We report the integration of this method within an ecological workflow management software, Predictive Ecosystem Analyzer (PEcAn), and its application and validation with two process-based terrestrial ecosystem models: SIPNET and ED2. In a test against a synthetic dataset, the emulator was able to retrieve the true parameter values. A comparison of the emulator approach to standard <q>brute-force</q> MCMC involving multiple data constraints showed that the emulator method was able to constrain the faster and simpler SIPNET model's parameters with comparable performance to the brute-force approach but reduced computation time by more than 2 orders of magnitude. The emulator was then applied to calibration of the ED2 model, whose complexity precludes standard (brute-force) Bayesian data assimilation techniques. Both models are constrained after assimilation of the observational data with the emulator method, reducing the uncertainty around their predictions. Performance metrics showed increased agreement between model predictions and data. Our study furthers efforts toward reducing model uncertainties, showing that the emulator method makes it possible to efficiently calibrate complex models.
Finley BK, Dijkstra P, Rasumussen C, Schwarz E, Mau RL, Liu XJ, van Gestel N, Hugate BA (2018) Soil mineral assemblage and substrate quality effects on microbial priming. Geoderma 322(38-47).
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Read PublicationNative soil organic carbon (SOC) decomposition rates may be altered through increased carbon (C) input, a phenomenon known as SOC priming (Blagodatskaya et al., 2011). Quantifying priming is important because it may modulate long-term SOC storage in ecosystems and therefore C biogeochemical cycling. Priming is positive when more SOC is decomposed or, conversely, negative when less native SOC is decomposed after C amendment (Kuzyakov et al., 2000; Kuzyakov, 2002; Bader and Cheng, 2007). Yet, controls over the direction and magnitude of the priming effect and the consequences for soil C balance remain uncertain (Dijkstra et al., 2013; Liu et al., 2017).
Forbes, WL, Mao J, Jin M, Kao SC, Fu W, Shi X, Riccuito DM, Thornton PE, Ribes A, Wang Y, Piao S, Zhao T, Schwalm CR, Hoffman FM, Fischer JB, Ito A, Poulter B, Fang Y, Tian H, Jain AK, Hayes DJ (2018) Contribution of environmental forcings to US runoff changes for the period 1950–2010. Environmental Research Letters 13(5): 054023.
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Read PublicationRunoff in the United States is changing, and this study finds that the measured change is dependent on the geographic region and varies seasonally. Specifically, observed annual total runoff had an insignificant increasing trend in the US between 1950 and 2010, but this insignificance was due to regional heterogeneity with both significant and insignificant increases in the eastern, northern, and southern US, and a greater significant decrease in the western US. Trends for seasonal mean runoff also differed across regions. By region, the season with the largest observed trend was autumn for the east (positive), spring for the north (positive), winter for the south (positive), winter for the west (negative), and autumn for the US as a whole (positive). Based on the detection and attribution analysis using gridded WaterWatch runoff observations along with semi-factorial land surface model simulations from the Multi-scale Synthesis and Terrestrial Model Intercomparison Project (MsTMIP), we found that while the roles of CO 2 concentration, nitrogen deposition, and land use and land cover were inconsistent regionally and seasonally, the effect of climatic variations was detected for all regions and seasons, and the change in runoff could be attributed to climate change in summer and autumn in the south and in autumn in the west. We also found that the climate-only and historical transient simulations consistently underestimated the runoff trends, possibly due to precipitation bias in the MsTMIP driver or within the models themselves.
Furze ME, Huggett BA, Aubrecth DM, Stolz CD, Carbone MS, Richardson AD (2018) Whole-tree nonstructural carbohydrate storage and seasonal dynamics in five temperate species. New Phytologist.
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Read PublicationDespite the importance of nonstructural carbohydrates (NSC) for growth and survival in woody plants, we know little about whole-tree NSC storage. The conventional theory suggests that NSC reserves will increase over the growing season and decrease over the dormant season. Here, we compare storage in five temperate tree species to determine the size and seasonal fluctuation of whole-tree total NSC pools as well as the contribution of individual organs. NSC concentrations in the branches, stemwood, and roots of 24 trees were measured across 12 months. We then scaled up concentrations to the whole-tree and ecosystem levels using allometric equations and forest stand inventory data. While whole-tree total NSC pools followed the conventional theory, sugar pools peaked in the dormant season and starch pools in the growing season. Seasonal depletion of total NSCs was minimal at the whole-tree level, but substantial at the organ level, particularly in branches. Surprisingly, roots were not the major storage organ as branches stored comparable amounts of starch throughout the year, and root reserves were not used to support springtime growth. Scaling up NSC concentrations to the ecosystem level, we find that commonly used, process-based ecosystem and land surface models all overpredict NSC storage.
Furze Morgan E, Jensen Ann M, Warren Jeffrey M, Richardson Andrew D (2018) Seasonal patterns of nonstructural carbohydrate reserves in four woody boreal species. The Journal of the Torrey Botanical Society 145(4).
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Read PublicationPlants store nonstructural carbohydrates (NSCs), such as sugars and starch, to use as carbon and energy sources for daily maintenance and growth needs as well as during times of stress. Allocation of NSCs to storage provides an important physiological strategy associated with future growth and survival, and thus understanding the seasonal patterns of NSC reserves provides insight into how species with different traits (e.g., growth form, leaf habit, wood anatomy) may respond to stress. We characterized the seasonal patterns of NSCs in four woody boreal plant species in Minnesota, USA. Sugar and starch concentrations were measured across the year in the roots and branches of two conifer trees, black spruce (Picea mariana (Mill.) B.S.P.) and eastern tamarack (Larix laricina (Du Roi) K. Koch), as well as in the leaves and branches of two evergreen broadleaf shrubs, bog Labrador tea (Rhododendron groenlandicum (Oeder) Kron & Judd) and leatherleaf (Chamaedaphne calyculata (L.) Moench). In general, seasonal variation was dominated by changes in starch across all organs and species. While similar seasonal patterns of NSCs were observed in the shrubs, different seasonal patterns were observed between the trees, particularly in the roots. Our results suggest that species-specific traits likely have consequences for organ-level storage dynamics, which may influence whole-plant growth and survival under global change.
Gibson CA, Koch BJ, Compson ZG, Hungate BA, Marks JC (2018) Ecosystem responses to restored flow in a travertine river. Freshwater Science 37(1): 169-177.
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Read PublicationDisruptions of natural flow impair rivers and streams worldwide. Those conducting restoration efforts have rarely explored how and when stream ecosystems can recover after reinstating natural flows. We quantified responses of ecosystem metabolism and N dynamics to the decommissioning and removal of a 100-y-old diversion dam in a desert stream, Fossil Creek, Arizona. Fossil Creek is a travertine river, meaning that CaCO3 concentrations in water in the springs that feed Fossil Creek are high enough to precipitate out of the water to form travertine terraces and deep pools. The majority of flow was diverted for power generation, so travertine deposition rates were significantly reduced and travertine terraces were smaller and less frequent compared to pre-dam historical records. Flow restoration enabled the recovery of the geochemical process of travertine deposition and increased gross primary production and N uptake to rates comparable to those measured in an upstream, reference reach. Reinstating a river’s natural flow regime can result in rapid and near-complete recovery of fundamental ecosystem processes that reshape the aquatic food web.
Granath G, Rydin H, Baltzer JL, Bengtsson F, Boncek N, Bragazza L, Bu ZJ, Caporn SJM, Dorrepaal E, Galanina O, Gałka M, Ganeva A, Gillikin DP, Goia I, Goncharova N, Hájek M, Haraguchi A, Harris LI, Humphreys E, Jiroušek M, Kajukało K, Karofeld E, Koronatova NG, Kosykh NP, Lamentowicz M, Lapshina E, Limpens J, Linkosalmi M, Ma JZ, Mauritz M, Munir TM, Natali SM, Natcheva R, Noskova M, Payne RJ, Pilkington K, Robinson S, Robroek BJM, Rochefort L, Singer D, Stenøien HK, Tuittila ES, Vellak K, Verheyden A, Waddington JM, Rice SK (2018) Environmental and taxonomic controls of carbon and oxygen stable isotope composition in Sphagnum across broad climatic and geographic ranges. Biogeosciences 16: 5189-5202.
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Read PublicationRain-fed peatlands are dominated by peat mosses (Sphagnum sp.), which for their growth depend on nutrients, water and CO2 uptake from the atmosphere. As the isotopic composition of carbon (12,13C) and oxygen (16,18O) of these Sphagnum mosses are affected by environmental conditions, Sphagnum tissue accumulated in peat constitutes a potential long-term archive that can be used for climate reconstruction. However, there is inadequate understanding of how isotope values are influenced by environmental conditions, which restricts their current use as environmental and palaeoenvironmental indicators. Here we tested (i) to what extent C and O isotopic variation in living tissue of Sphagnum is species-specific and associated with local hydrological gradients, climatic gradients (evapotranspiration, temperature, precipitation) and elevation; (ii) whether the C isotopic signature can be a proxy for net primary productivity (NPP) of Sphagnum; and (iii) to what extent Sphagnum tissue δ18O tracks the δ18O isotope signature of precipitation. In total, we analysed 337 samples from 93 sites across North America and Eurasia using two important peat-forming Sphagnum species (S. magellanicum, S. fuscum) common to the Holarctic realm. There were differences in δ13C values between species. For S. magellanicum δ13C decreased with increasing height above the water table (HWT, R2 = 17%) and was positively correlated to productivity (R2 = 7%). Together these two variables explained 46% of the between-site variation in δ13C values. For S. fuscum, productivity was the only significant predictor of δ13C but had low explanatory power (total R2 = 6%). For δ18O values, approximately 90% of the variation was found between sites. Globally modelled annual δ18O values in precipitation explained 69% of the between-site variation in tissue δ18O. S. magellanicum showed lower δ18O enrichment than S. fuscum (−0.83‰ lower). Elevation and climatic variables were weak predictors of tissue δ18O values after controlling for δ18O values of the precipitation. To summarize, our study provides evidence for (a) good predictability of tissue δ18O values from modelled annual δ18O values in precipitation, and (b) the possibility of relating tissue δ13C values to HWT and NPP, but this appears to be species-dependent. These results suggest that isotope composition can be used on a large scale for climatic reconstructions but that such models should be species-specific.
Guo JS, Hungate BA, Kolb TE, Koch GW (2018) Water source niche overlap increases with site moisture availability in woody perennials. Plant Ecology.
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Read PublicationClassical niche partitioning theory posits increased competition for and partitioning of the most limiting resource among coexisting species. Coexisting plant species may vary in rooting depth, reflecting niche partitioning in water source use. Our goal was to assess the soil water partitioning of woody plant communities across northern Arizona along an elevational moisture gradient using stem and soil water isotopes from two sampling periods to estimate the use of different water sources. We hypothesized that niche overlap of water sources would be higher and monsoon precipitation uptake would be lower at sites with higher moisture availability. Pairwise niche overlap of coexisting species was calculated using mixing model estimates of proportional water use for three sources. Across the moisture gradient, niche overlap increased with site moisture index (precipitation/potential evapotranspiration) across seasons, and site moisture index explained 37% of the variation in niche overlap of intermediate and deeper sources of water. Desert trees utilized more winter source water than desert shrubs, suggesting the partitioning of water sources between functional groups. However, seasonal differences in surface water use were primarily found at intermediate levels of site moisture availability. Our findings support classical niche partitioning theory in that plants exhibit higher overlap of water sources when water is not a limiting resource.
Guo X, Feng J, Shi Z, Zhou X, Yuan M, Tao X, Hale L, Yuan T, Wang J, Qin Y, Zhou A, Fu Y, Wu L, He Z, Van Nostrand JD, Ning D, Liu X, Luo Y, Tiedje JM, Yang Y, Zhou J (2018) Climate warming leads to divergent succession of grassland microbial communities. Nature Climate Change 8(9): 813-818.
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Read PublicationAccurate climate projections require an understanding of the effects of warming on ecological communities and the underlying mechanisms that drive them1–3. However, little is known about the effects of climate warming on the succession of microbial communities4,5. Here we examined the temporal succession of soil microbes in a long-term climate change experiment at a tall-grass prairie ecosystem. Experimental warming was found to significantly alter the community structure of bacteria and fungi. By determining the time-decay relationships and the paired differences of microbial communities under warming and ambient conditions, experimental warming was shown to lead to increasingly divergent succession of the soil microbial communities, with possibly higher impacts on fungi than bacteria. Variation partition- and null model-based analyses indicate that stochastic processes played larger roles than deterministic ones in explaining microbial community taxonomic and phylogenetic compositions. However, in warmed soils, the relative importance of stochastic processes decreased over time, indicating a potential deterministic environmental filtering elicited by warming. Although successional trajectories of microbial communities are difficult to predict under future climate change scenarios, their composition and structure are projected to be less variable due to warming-driven selection.
He W, Ju W, Schwalm CR, Sippel S, Wu X, He Q, Song L, Zhang C, Li J, Sitch S, Viovy N, Friedlingstein P, Jain AK (2018) Large-scale droughts responsible for dramatic reductions of terrestrial net carbon uptake over North America in 2011 and 2012. Journal of Geophysical Research: Biogeosciences 123(7): 2053-2071.
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Read PublicationRecently, severe droughts that occurred in North America are likely to have impacted its terrestrial carbon sink. However, process-based understanding of how meteorological conditions prior to the onset of drought, for instance warm or cold springs, affect drought-induced carbon cycle effects remains scarce. Here we assess and compare the response of terrestrial carbon fluxes to summer droughts in 2011 and 2012 characterized by contrasting spring conditions. The analysis is based on a comprehensive ensemble of carbon cycle models, including FLUXCOM, TRENDY v5, SiBCASA, CarbonTracker Europe, and CarbonTracker, and emerging Earth observations. In 2011, large reductions of net ecosystem production (NEP; ?0.24 ± 0.17 Pg C/year) are due to decreased gross primary production (?0.17 ± 0.18 Pg C/year) and slightly increased ecosystem respiration (+0.07 ± 0.17 Pg C/year). Conversely, in 2012, NEP reductions (?0.17 ± 0.25 Pg C/year) are attributed to a larger increase of ecosystem respiration (+0.48 ± 0.27 Pg C/year) than gross primary production (+0.31 ± 0.29 Pg C/year), induced predominantly by an extra warmer spring prior to summer drought. Two temperate ecoregions crops/agriculture and the grass/shrubs contribute largest to these reductions and also dominate the interannual variations of NEP during 2007?2014. Moreover, the warming spring compensated largely the negative carbon anomaly due to summer drought, consistent with earlier studies; however, the compensation occurred only in some specific ecoregions. Overall, our analysis offers a refined view on recent carbon cycle variability and extremes in North America. It corroborates earlier results but also highlights differences with respect to ecoregion-specific carbon cycle responses to drought and heat.
Hemming DL, Abernethy R, Armitage C, Bolmgren K, Myneni R, Park T, Richardson AD, Rutishauser T, Sparks TH, Thakeray SJ (2018) Sidebar 2.3. Phenology of terrestrial and freshwater primary producers. Bulletin of the American Meteorological Society.
Read AbstractPhenology is the study of recurring events in nature and their relationships with climate. The word derives from the Greek phaínō ‘appear’ and logos ‘reason’, emphasizing the focus on observing events and understanding why they occur (Demarée and Rutishauser 2009). Phenological recording has a history that dates back many centuries (Linneaus and Bark 1753; Aono and Kazui 2008). More recently, advances in monitoring technologies have enabled automated and remotely sensed observations, complemented by increasing citizen science participation in monitoring efforts. Phenological information can also be derived from widespread environmental monitoring stations around the globe.
Phenological records clearly demonstrate the biological effects of year-to-year variability in climate, as well as longer-term trends associated with environmental change. Phenological monitoring thus plays an important role in understanding how our planet is changing. Changes in the growing season, for example, are more tangible and more readily conveyed to the general public than seemingly small changes in mean annual temperature.
Here, we describe just a fraction of the phenological information currently available, highlighting northern hemisphere records of phenology of primary producers across a range of spatial and temporal scales.
Hewitt RE, Taylor DL, Genet H, McGuire AD, Mack MC (2018) Below-ground plant traits influence tundra plant acquisition of newly thawed permafrost nitrogen. Journal of Ecology.
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Read PublicationThe release of permafrost-derived nitrogen (N) has the potential to fertilize tundra vegetation, which in turn may stimulate productivity and thus offset carbon (C) losses from thawing permafrost. Below-ground plant traits may mediate ecosystem response to permafrost thaw and associated feedbacks to the atmosphere by differentially conferring access to deep, newly thawed permafrost N. Yet, identifying roots and quantifying root N uptake from deep, cold soils in complex plant communities has proved challenging to date. We investigated plant acquisition of experimentally added 15N isotope tracer applied at the permafrost boundary in graminoid- and shrub-dominated tundra at Eight Mile Lake, Alaska, when the thaw front was close to its maximum depth, simulating the release of newly thawed permafrost N. We used molecular tools to verify species and estimate biomass, nitrogen, and isotope pools. Root biomass depth distributions follow an asymptotic relationship with depth, typical of other ecosystems. Few species had roots occurring close to the thaw front. Rubus chamaemorus, a short-statured non-mycorrhizal forb, and Carex bigelowii, a sedge, consistently had the deepest roots. Twenty-four hours after isotope addition, we observed that deep-rooted, non-mycorrhizal species had the highest 15N enrichment values in their fine root tissue indicating that they access deep N late in the growing season when the thaw front is deepest. Deep-rooted plants are therefore able to immediately take up newly thawed permafrost-derived N. During the following growing season, herbaceous, non-mycorrhizal plants allocated tracer above-ground before woody, mycorrhizal plants. Ectomycorrhizal deciduous and ericoid mycorrhizal evergreen shrubs, by contrast, did not have immediate access to the deep N tracer and assimilated it into new foliar tissue gradually over the following growing season. Synthesis. Graminoids and forbs that have immediate access to deep N represent a modest C sink compared to C emissions from thawing permafrost. However, the effects of deep N fertilization on shrubs over longer time-scales may stimulate productivity and account for a more considerable N and C sink, thus constraining the permafrost C-climate feedback.
Holdo RM, Nippert JB, Mack MC (2018) Rooting depth varies differentially in trees and grasses as a function of mean annual rainfall in an African savanna. Oecologia 186(1): 269-280.
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Read PublicationA significant fraction of the terrestrial biosphere comprises biomes containing tree–grass mixtures. Forecasting vegetation dynamics in these environments requires a thorough understanding of how trees and grasses use and compete for key belowground resources. There is disagreement about the extent to which tree–grass vertical root separation occurs in these ecosystems, how this overlap varies across large-scale environmental gradients, and what these rooting differences imply for water resource availability and tree–grass competition and coexistence. To assess the extent of tree–grass rooting overlap and how tree and grass rooting patterns vary across resource gradients, we examined landscape-level patterns of tree and grass functional rooting depth along a mean annual precipitation (MAP) gradient extending from ~ 450 to ~ 750 mm year−1 in Kruger National Park, South Africa. We used stable isotopes from soil and stem water to make inferences about relative differences in rooting depth between these two functional groups. We found clear differences in rooting depth between grasses and trees across the MAP gradient, with grasses generally exhibiting shallower rooting profiles than trees. We also found that trees tended to become more shallow-rooted as a function of MAP, to the point that trees and grasses largely overlapped in terms of rooting depth at the wettest sites. Our results reconcile previously conflicting evidence for rooting overlap in this system, and have important implications for understanding tree–grass dynamics under altered precipitation scenarios.
Holland-Moritz H, Stuart J, Lewis LR, Miller S, Mack MC, McDaniel SF, Fierer N (2018) Novel bacterial lineages associated with boreal moss species. Environmental Microbiology 20(7):2625-2638.
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Read PublicationMosses are critical components of boreal ecosystems where they typically account for a large proportion of net primary productivity and harbour diverse bacterial communities that can be the major source of biologically-fixed nitrogen in these ecosystems. Despite their ecological importance, we have limited understanding of how microbial communities vary across boreal moss species and the extent to which local site conditions may influence the composition of these bacterial communities. We used marker gene sequencing to analyze bacterial communities associated with seven boreal moss species collected near Fairbanks, AK, USA. We found that host identity was more important than site in determining bacterial community composition and that mosses harbour diverse lineages of potential N2-fixers as well as an abundance of novel taxa assigned to understudied bacterial phyla (including candidate phylum WPS-2). We performed shotgun metagenomic sequencing to assemble genomes from the WPS-2 candidate phylum and found that these moss-associated bacteria are likely anoxygenic phototrophs capable of carbon fixation via RuBisCo with an ability to utilize byproducts of photorespiration from hosts via a glyoxylate shunt. These results give new insights into the metabolic capabilities of understudied bacterial lineages that associate with mosses and the importance of plant hosts in shaping their microbiomes.
Hou E, Chen C, Luo Y, Zhou G, Kuang Y, Zhang Y, Heenan M, Lu X, Wen D (2018) Effects of climate on soil phosphorus cycle and availability in natural terrestrial ecosystems. Global Change Biology.
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Read PublicationClimate is predicted to change over the 21st century. However, little is known about how climate change can affect soil phosphorus (P) cycle and availability in global terrestrial ecosystems, where P is a key limiting nutrient. With a global database of Hedley P fractions and key‐associated physiochemical properties of 760 (seminatural) natural soils compiled from 96 published studies, this study evaluated how climate pattern affected soil P cycle and availability in global terrestrial ecosystems. Overall, soil available P, indexed by Hedley labile inorganic P fraction, significantly decreased with increasing mean annual temperature (MAT) and precipitation (MAP). Hypothesis‐oriented path model analysis suggests that MAT negatively affected soil available P mainly by decreasing soil organic P and primary mineral P and increasing soil sand content. MAP negatively affected soil available P both directly and indirectly through decreasing soil primary mineral P; however, these negative effects were offset by the positive effects of MAP on soil organic P and fine soil particles, resulting in a relatively minor total MAP effect on soil available P. As aridity degree was mainly determined by MAP, aridity also had a relatively minor total effect on soil available P. These global patterns generally hold true irrespective of soil depth (less than or equal to 10 cm or greater than 10 cm) or site aridity index (less than or equal to 1.0 or greater than 1.0), and were also true for the low‐sand (less than or equal to 50%) soils. In contrast, available P of the high‐sand (greater than 50%) soils was positively affected by MAT and aridity and negatively affected by MAP. Our results suggest that temperature and precipitation have contrasting effects on soil P availability and can interact with soil particle size to control soil P availability.
Huang K, Xia J, Wang Y, Ahlstrom A, Chen J, Cook RB, Cui E, Fang Y, Fisher JB, Huntzinger DN, Li Z, Michalak AM, Qiao Y, Schaefer K, Schwalm C, Wang J, Wei Y, Xu X, Yan L, Bian C, Luo Y (2018) Enhanced peak growth of global vegetation and its key mechanisms. Nature Ecology & Evolution.
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Read PublicationThe annual peak growth of vegetation is critical in characterizing the capacity of terrestrial ecosystem productivity and shaping the seasonality of atmospheric CO2 concentrations. The recent greening of global lands suggests an increasing trend of terrestrial vegetation growth, but whether or not the peak growth has been globally enhanced still remains unclear. Here, we use two global datasets of gross primary productivity (GPP) and a satellite-derived Normalized Difference Vegetation Index (NDVI) to characterize recent changes in annual peak vegetation growth (that is, GPPmax and NDVImax). We demonstrate that the peak in the growth of global vegetation has been linearly increasing during the past three decades. About 65% of the NDVImax variation is evenly explained by expanding croplands (21%), rising CO2 (22%) and intensifying nitrogen deposition (22%). The contribution of expanding croplands to the peak growth trend is substantiated by measurements from eddy-flux towers, sun-induced chlorophyll fluorescence and a global database of plant traits, all of which demonstrate that croplands have a higher photosynthetic capacity than other vegetation types. The large contribution of CO2 is also supported by a meta-analysis of 466 manipulative experiments and 15 terrestrial biosphere models. Furthermore, we show that the contribution of GPPmax to the change in annual GPP is less in the tropics than in other regions. These multiple lines of evidence reveal an increasing trend in the peak growth of global vegetation. The findings highlight the important roles of agricultural intensification and atmospheric changes in reshaping the seasonality of global vegetation growth.
Huang Y, Lu X, Shi Z, Lawrence D, Koven CD, Xia J, Du Z, Kluzek E, Luo Y (2018) Matrix approach to land carbon cycle modeling: A case study with the Community Land Model. Global Change Biology 24(3): 1394-1404.
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Read PublicationThe terrestrial carbon (C) cycle has been commonly represented by a series of C balance equations to track C influxes into and effluxes out of individual pools in earth system models (ESMs). This representation matches our understanding of C cycle processes well but makes it difficult to track model behaviors. It is also computationally expensive, limiting the ability to conduct comprehensive parametric sensitivity analyses. To overcome these challenges, we have developed a matrix approach, which reorganizes the C balance equations in the original ESM into one matrix equation without changing any modeled C cycle processes and mechanisms. We applied the matrix approach to the Community Land Model (CLM4.5) with vertically‐resolved biogeochemistry. The matrix equation exactly reproduces litter and soil organic carbon (SOC) dynamics of the standard CLM4.5 across different spatial‐temporal scales. The matrix approach enables effective diagnosis of system properties such as C residence time and attribution of global change impacts to relevant processes. We illustrated, for example, the impacts of CO2 fertilization on litter and SOC dynamics can be easily decomposed into the relative contributions from C input, allocation of external C into different C pools, nitrogen regulation, altered soil environmental conditions, and vertical mixing along the soil profile. In addition, the matrix tool can accelerate model spin‐up, permit thorough parametric sensitivity tests, enable pool‐based data assimilation, and facilitate tracking and benchmarking of model behaviors. Overall, the matrix approach can make a broad range of future modeling activities more efficient and effective.
Hufkens K, Basler D, Milliman T, Melaas EK, Richardson AD (2018) An integrated phenology modelling framework in R. Methods in Ecology and Evolution 9(5): 1276-1285.
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Read PublicationAbstract Phenology is a first-order control on productivity and mediates the biophysical environment by altering albedo, surface roughness length and evapotranspiration. Accurate and transparent modelling of vegetation phenology is therefore key in understanding feedbacks between the biosphere and the climate system. Here, we present the PHENOR R package and modelling framework. The framework leverages measurements of vegetation phenology from four common phenology observation datasets, the PhenoCam network, the USA National Phenology Network (USA-NPN), the Pan European Phenology Project (PEP725), MODIS phenology (MCD12Q2) combined with (global) retrospective and projected climate data. We show an example analysis, using the phenor modelling framework, which quickly and easily compares 20 included spring phenology models for three plant functional types. An analysis of model skill using the root mean squared (RMSE) error shows little or no difference regardless of model structure, corroborating previous studies. We argue that addressing this issue will require novel model development combined with easy data assimilation as facilitated by our framework. In conclusion, we hope the phenor phenology modelling framework in the r language and environment for statistical computing will facilitate reproducibility and community driven phenology model development, in order to increase their overall predictive power, and leverage an ever growing number of phenology data products.
Jean M, Mack MC, Johnstone JF (2018) Spatial and temporal variation in moss-associated dinitrogen fixation in coniferous- and deciduous-dominated Alaskan boreal forests. Plant Ecology 219(7), 837-851.
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Read PublicationDominant canopy tree species have strong effects on the composition and function of understory species, particularly bryophytes. In boreal forests, bryophytes and their associated microbes are a primary source of ecosystem nitrogen (N) inputs, and an important process regulating ecosystem productivity. We investigated how feather moss-associated N<sub>2</sub>-fixation rates and contribution to N budgets vary in time and space among coniferous and broadleaf deciduous forests. We measured N<sub>2</sub>-fixation rates using stable isotope (<sup>15</sup>N<sub>2</sub>) labeling in two moss species (<em class="EmphasisTypeItalic ">Pleurozium schreberi</em> and <em class="EmphasisTypeItalic ">Hylocomium splendens</em>) in broadleaf deciduous (Alaska paper birch—<em class="EmphasisTypeItalic ">Betula neoalaskana</em>) and coniferous (black spruce—<em class="EmphasisTypeItalic ">Picea mariana</em>) stands near Fairbanks, interior Alaska, from 2013 to 2015. N<sub>2</sub>-fixation rates showed substantial inter-annual variation among the 3 years. High N<sub>2</sub>-fixation was more strongly associated with high precipitation than air temperature or light availability. Overall, contribution of N<sub>2</sub>-fixation to N budgets was greater in spruce than in birch stands. Our results enhance the knowledge of the processes that drive N<sub>2</sub>-fixation in boreal forests, which is important for predicting ecosystem consequences of changing forest composition.
Jeong SJ, Bloom AA, Schimel D, Sweeney C, Parazoo NC, Medvigy D, Schaepman-Strub G, Zheng C, Schwalm CR, Huntzinger DN, Michalak AM, Miller CE (2018) Accelerating rates of Arctic carbon cycling revealed by long-term atmospheric CO2 measurements. Science Advances 4(7): eaao1167.
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Read PublicationThe contemporary Arctic carbon balance is uncertain, and the potential for a permafrost carbon feedback of anywhere from 50 to 200 petagrams of carbon (Schuur et al., 2015) compromises accurate 21st-century global climate system projections. The 42-year record of atmospheric CO2 measurements at Barrow, Alaska (71.29 N, 156.79 W), reveals significant trends in regional land-surface CO2 anomalies (ΔCO2), indicating long-term changes in seasonal carbon uptake and respiration. Using a carbon balance model constrained by ΔCO2, we find a 13.4% decrease in mean carbon residence time (50% confidence range = 9.2 to 17.6%) in North Slope tundra ecosystems during the past four decades, suggesting a transition toward a boreal carbon cycling regime. Temperature dependencies of respiration and carbon uptake suggest that increases in cold season Arctic labile carbon release will likely continue to exceed increases in net growing season carbon uptake under continued warming trends.
Jiang J, Haung Y, Ma S, Stacy M, Shi Z, Ricciuto DM, Hanson PJ, Luo Y (2018) Forecasting responses of a northern peatland carbon cycle to elevated CO2 and a gradient of experimental warming. Journal of Geophysical Research: Biogeosciences.
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Read PublicationThe ability to forecast ecological carbon cycling is imperative to land management in a world where past carbon fluxes are no longer a clear guide in the Anthropocene. However, carbon‐flux forecasting has not been practiced routinely like numerical weather prediction. This study explored (1) the relative contributions of model forcing data and parameters to uncertainty in forecasting flux‐ versus pool‐based carbon cycle variables and (2) the time points when temperature and CO2 treatments may cause statistically detectable differences in those variables. We developed an online forecasting workflow (Ecological Platform for Assimilation of Data (EcoPAD)), which facilitates iterative data‐model integration. EcoPAD automates data transfer from sensor networks, data assimilation, and ecological forecasting. We used the Spruce and Peatland Responses Under Changing Experiments data collected from 2011 to 2014 to constrain the parameters in the Terrestrial Ecosystem Model, forecast carbon cycle responses to elevated CO2 and a gradient of warming from 2015 to 2024, and specify uncertainties in the model output. Our results showed that data assimilation substantially reduces forecasting uncertainties. Interestingly, we found that the stochasticity of future external forcing contributed more to the uncertainty of forecasting future dynamics of C flux‐related variables than model parameters. However, the parameter uncertainty primarily contributes to the uncertainty in forecasting C pool‐related response variables. Given the uncertainties in forecasting carbon fluxes and pools, our analysis showed that statistically different responses of fast‐turnover pools to various CO2 and warming treatments were observed sooner than slow‐turnover pools. Our study has identified the sources of uncertainties in model prediction and thus leads to improve ecological carbon cycling forecasts in the future.
Klosterman S, Hufkens K, Richardson AD (2018) Later springs green-up faster: The relation between onset and completion of green-up in deciduous forests of North America. International Journal of Biometeorology 62(9): 1645-1655.
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Read PublicationIn deciduous forests, spring leaf phenology controls the onset of numerous ecosystem functions. While most studies have focused on a single annual spring event, such as budburst, ecosystem functions like photosynthesis and transpiration increase gradually after budburst, as leaves grow to their mature size. Here, we examine the “velocity of green-up,” or duration between budburst and leaf maturity, in deciduous forest ecosystems of eastern North America. We use a diverse data set that includes 301 site-years of phenocam data across a range of sites, as well as 22 years of direct ground observations of individual trees and 3 years of fine-scale high-frequency aerial photography, both from Harvard Forest. We find a significant association between later start of spring and faster green-up: − 0.47 ± 0.04 (slope ± 1 SE) days change in length of green-up for every day later start of spring within phenocam sites, − 0.31 ± 0.06 days/day for trees under direct observation, and − 1.61 ± 0.08 days/day spatially across fine-scale landscape units. To explore the climatic drivers of spring leaf development, we fit degree-day models to the observational data from Harvard Forest. We find that the default phenology parameters of the ecosystem model PnET make biased predictions of leaf initiation (39 days early) and maturity (13 days late) for red oak, while the optimized model has biases of 1 day or less. Springtime productivity predictions using optimized parameters are closer to results driven by observational data (within 1%) than those of the default parameterization (17% difference). Our study advances empirical understanding of the link between early and late spring phenophases and demonstrates that accurately modeling these transitions is important for simulating seasonal variation in ecosystem productivity.
Klosterman S, Melaas E, Wang J, Martinez A, Frederick S, O'Keefe J, Orwig DA, Wang Z, Sun Q, Schaaf C, Friedl M, Richardson AD (2018) Fine-scale perspectives on landscape phenology from unmanned aerial vehicle (UAV) photography. Agricultural and Forest Meteorology 248: 397-407.
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Read PublicationForest phenology is a multi-scale phenomenon, arising from processes in leaves and trees, with effects on the ecology of plant communities and landscapes. Because phenology controls carbon and water cycles, which are commonly observed at the ecosystem scale (e.g. eddy flux measurements), it is important to characterize the relation between phenophase transition events at different spatial scales. We use aerial photography recorded from an unmanned aerial vehicle (UAV) to observe plant phenology over a large area (5.4 ha) and across diverse communities, with spatial and temporal resolution at the scale of individual tree crowns and their phenophase transition events (10 m spatial resolution, ∼5 day temporal resolution in spring, weekly in autumn). We validate UAV-derived phenophase transition dates through comparison with direct observations of tree phenology, PhenoCam image analysis, and satellite remote sensing. We then examine the biological correlates of spatial variance in phenology using a detailed species inventory and land cover classification. Our results show that species distribution is the dominant factor in spatial variability of ecosystem phenology. We also explore statistical relations governing the scaling of phenology from an organismic scale (10 m) to forested landscapes (1 km) by analyzing UAV photography alongside Landsat and MODIS data. From this analysis we find that spatial standard deviation in transition dates decreases linearly with the logarithm of increasing pixel size. We also find that fine-scale phenology aggregates to a coarser scale as the median and not the mean date in autumn, indicating coarser scale phenology is less sensitive to the tails of the distribution of sub-pixel transitions in the study area. Our study is the first to observe forest phenology in a spatially comprehensive, whole-ecosystem way, yet with fine enough spatial resolution to describe organism-level correlates and scaling phenomena.
Koch BJ, McHugh TA, Hayer M, Schwartz E, Blazewicz SJ, Dijkstra P, van Gestel N, Marks JC, Mau RL, Morrissey EM, Pett-Ridge J, Hungate BA (2018) Estimating taxon-specific population dynamics in diverse microbial communities. Ecosphere 9(1):e02090.
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Read PublicationUnderstanding how population-level dynamics contribute to ecosystem-level processes is a primary focus of ecological research and has led to important breakthroughs in the ecology of macroscopic organisms. However, the inability to measure population-specific rates, such as growth, for microbial taxa within natural assemblages has limited ecologists’ understanding of how microbial populations interact to regulate ecosystem processes. Here, we use isotope incorporation within DNA molecules to model taxon-specific population growth in the presence of 18O-labeled water. By applying this model to phylogenetic marker sequencing data collected from stable-isotope probing studies, we estimate rates of growth, mortality, and turnover for individual microbial populations within soil assemblages. When summed across the entire bacterial community, our taxon-specific estimates are within the range of other whole-assemblage measurements of bacterial turnover. Because it can be applied to environmental samples, the approach we present is broadly applicable to measuring population growth, mortality, and associated biogeochemical process rates of microbial taxa for a wide range of ecosystems and can help reveal how individual microbial populations drive biogeochemical fluxes.
Kosmala M, Hufkens K, Richardson AD (2018) Integrating camera imagery, crowdsourcing, and deep learning to improve high-frequency automated monitoring of snow at continental-to-global scales. PLOS ONE 13(12): e0209649.
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Read PublicationSnow is important for local to global climate and surface hydrology, but spatial and temporal heterogeneity in the extent of snow cover make accurate, fine-scale mapping and monitoring of snow an enormous challenge. We took 184,453 daily near-surface images acquired by 133 automated cameras and processed them using crowdsourcing and deep learning to determine whether snow was present or absent in each image. We found that the crowdsourced data had an accuracy of 99.1% when compared with expert evaluation of the same imagery. We then used the image classification to train a deep convolutional neural network via transfer learning, with accuracies of 92% to 98%, depending on the image set and training method. The majority of neural network errors were due to snow that was present not being detected. We used the results of the neural networks to validate the presence or absence of snow inferred from the MODIS satellite sensor and obtained similar results to those from other validation studies. This method of using automated sensors, crowdsourcing, and deep learning in combination produced an accurate high temporal dataset of snow presence across a continent. It holds broad potential for real-time large-scale acquisition and processing of ecological and environmental data in support of monitoring, management, and research objectives.
Landhausser SM, Chow PS, Adams HD, Dickman LT, Furze ME, Richardson AD, Gleixner G, Hartmann H, Kuhlman I, Hoch G, Schmid S, Richter A, Wiesenbauer J, Wild B, McDowell NG (2018) Standardized protocols and procedures can precisely and accurately quantify non-structural carbohydrates. Tree Physiology 38(12):1764-1778.
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Read PublicationNon-structural carbohydrates (NSCs), the stored products of photosynthesis, building blocks for growth and fuel for respiration, are central to plant metabolism, but their measurement is challenging. Differences in methods and procedures among laboratories can cause results to vary widely, limiting our ability to integrate and generalize patterns in plant carbon balance among studies. A recent assessment found that NSC concentrations measured for a common set of samples can vary by an order of magnitude, but sources for this variability were unclear. We measured a common set of nine plant material types, and two synthetic samples with known NSC concentrations, using a common protocol for sugar extraction and starch digestion, and three different sugar quantification methods (ion chromatography, enzyme, acid) in six laboratories. We also tested how sample handling, extraction solvent and centralizing parts of the procedure in one laboratory affected results. Non-structural carbohydrate concentrations measured for synthetic samples were within about 11.5% of known values for all three methods. However, differences among quantification methods were the largest source of variation in NSC measurements for natural plant samples because the three methods quantify different NSCs. The enzyme method quantified only glucose, fructose and sucrose, with ion chromatography we additionally quantified galactose, while the acid method quantified a large range of mono- and oligosaccharides. For some natural samples, sugars quantified with the acid method were two to five times higher than with other methods, demonstrating that trees allocate carbon to a range of sugar molecules. Sample handling had little effect on measurements, while ethanol sugar extraction improved accuracy over water extraction. Our results demonstrate that reasonable accuracy of NSC measurements can be achieved when different methods are used, as long as protocols are robust and standardized. Thus, we provide detailed protocols for the extraction, digestion and quantification of NSCs in plant samples, which should improve the comparability of NSC measurements among laboratories.
Lee MS, Hollinger DY, Keenan TF, Ouimette AP, Ollinger SV, Richardson AD (2018) Model-based analysis of the impact of diffuse radiation on CO2 exchange in a temperate deciduous forest. Agricultural and Forest Meteorology 249: 377-389.
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Read PublicationClouds and aerosols increase the fraction of global solar irradiance that is diffuse light. This phenomenon is known to increase the photosynthetic light use efficiency (LUE) of closed-canopy vegetation by redistributing photosynthetic photon flux density (400–700 nm) from saturated, sunlit leaves at the top of the canopy, to shaded leaves deeper in the canopy. We combined a process-based carbon cycle model with 10 years of eddy covariance carbon flux measurements and other ancillary data sets to assess 1) how this LUE enhancement influences interannual variation in carbon uptake, and 2) how errors in modeling diffuse fraction affect predictions of carbon uptake. Modeled annual gross primary productivity (GPP) increased by ≈0.94% when observed levels of diffuse fraction were increased by 0.01 (holding total irradiance constant). The sensitivity of GPP to increases in diffuse fraction was highest when the diffuse fraction was low to begin with, and lowest when the diffuse fraction was already high. Diffuse fraction also explained significantly more of the interannual variability of modeled net ecosystem exchange (NEE), than did total irradiance. Two tested radiation partitioning models yielded over- and underestimates of diffuse fraction at our site, which propagated to over- and underestimates of annual NEE, respectively. Our findings highlight the importance of incorporating LUE enhancement under diffuse light into models of global primary production, and improving models of diffuse fraction.
Lei L, Xia J, Li X, Huang K, Zhang A, Chen S, Weng E, Luo Y, Wan S (2018) Water response of ecosystem respiration regulates future projection of net ecosystem productivity in a semiarid grassland. Agricultural and Forest Meteorology (252) 175-191.
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Read PublicationRecent evidences show that terrestrial biogeochemical models have large uncertainty in estimating climate-change effect on grassland net ecosystem productivity (NEP), which is defined as the difference between gross ecosystem photosynthesis (GEP) and respiration (ER). It remains unclear that whether GEP or ER limits the model capability to simulate NEP responses to climate change in semiarid grasslands. Given the surrogate CENTURY-type model is widely used for Earth system modeling, we investigated two of them (i.e., DAYCENT and TECO models) and examined which processes dominate their ability to capture the responses of NEP to experimental climate changes in a temperate steppe of northern China. During the simulation from 2006 to 2008, the two models captured the observed mean annual NEP in the control plots when they were validated by the observations from an adjacent eddy-flux tower. However, they failed to capture the treatment effects of experimental warming and increased precipitation on NEP because of the poor estimations of ER responses. DAYCENT model simulated a higher precipitation effect on ER (37.83%) and TECO model overestimated the warming effect on ER by 8.18%. The simulation of treatment effects on ER and therefore NEP can be improved by an optimized parameterization of the water-related decay functions for soil organic carbon (C). The simulated cumulative loss of total ecosystem C stock during 2010–2100 were decreased when the TECO model used experiment-fitted parameters (0.72 kg C m−2) instead of using the initial validation with eddy-flux data (0.96 kg C m−2). The ecosystem shifted from C sink to source at threshold of 435 mm of annual total precipitation. Our findings indicate that future projection of C cycle in semiarid grasslands could be improved by better understanding of water response of ecosystem respiratory processes.
Levy RC, Burakowski E, Richardson AD (2018) Novel measurements of a fine-scale albedo: Using a commercial quadcopter to measure radiation fluxes. Remote Sensing 10(8).
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Download .PDFRemote sensing of radiative indices must balance spatially and temporally coarse satellite measurements with finer-scale, but geographically limited, in-situ surface measurements. Instruments mounted upon an Unmanned Aerial Vehicle (UAV) can provide small-scale, mobile remote measurements that fill this resolution gap. Here we present and validate a novel method of obtaining albedo values using an unmodified quadcopter at a deciduous northern hardwood forest. We validate this method by comparing simultaneous albedo estimates by UAV and a fixed tower at the same site. We found that UAV provided stable albedo measurements across multiple flights, with results that were well within the range of tower-estimated albedo at similar forested sites. Our results indicate that in-situ albedo measurements (tower and UAV) capture more site-to-site variation in albedo than satellite measurements. Overall, we show that UAVs produce reliable, consistent albedo measurements that can capture crucial surface heterogeneity, clearly distinguishing between different land uses. Future application of this approach can provide detailed measurements of albedo and potentially other vegetation indices to enhance global research and modeling efforts.
Li Q, Lu X, Wang Y, Huang X, Cox PM, Luo Y (2018) Leaf area index identified as a major source of variability in modeled CO2 fertilization. Biogeosciences 15: 6909-6925.
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Read PublicationThe concentration–carbon feedback (β), also called the CO2 fertilization effect, is a key unknown in climate–carbon-cycle projections. A better understanding of model mechanisms that govern terrestrial ecosystem responses to elevated CO2 is urgently needed to enable a more accurate prediction of future terrestrial carbon sink. We conducted C-only, carbon–nitrogen (C–N) and carbon–nitrogen–phosphorus (C–N–P) simulations of the Community Atmosphere Biosphere Land Exchange model (CABLE) from 1901 to 2100 with fixed climate to identify the most critical model process that causes divergence in β. We calculated CO2 fertilization effects at various hierarchical levels from leaf biochemical reaction and leaf photosynthesis to canopy gross primary production (GPP), net primary production (NPP), and ecosystem carbon storage (cpool) for seven C3 plant functional types (PFTs) in response to increasing CO2 under the RCP 8.5 scenario. Our results show that β values at biochemical and leaf photosynthesis levels vary little across the seven PFTs, but greatly diverge at canopy and ecosystem levels in all simulations. The low variation of the leaf-level β is consistent with a theoretical analysis that leaf photosynthetic sensitivity to increasing CO2 concentration is almost an invariant function. In the CABLE model, the major jump in variation of β values from leaf levels to canopy and ecosystem levels results from divergence in modeled leaf area index (LAI) within and among PFTs. The correlation of βGPP, βNPP, or βcpool each with βLAI is very high in all simulations. Overall, our results indicate that modeled LAI is a key factor causing the divergence in β in the CABLE model. It is therefore urgent to constrain processes that regulate LAI dynamics in order to better represent the response of ecosystem productivity to increasing CO2 in Earth system models.
Liang G, Cai A, Wu H, Wu X, Houssou AA, Ren C, Wang Z, Gao L, Wang B, Li S, Song X, Cai D (2018) Soil biochemical parameters in the rhizosphere contribute more to changes in soil respiration and its components than those in the bulk soil under nitrogen application in croplands. Plant and Soil.
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Read PublicationSoil respiration (RS), which is the second largest carbon flux between the atmosphere and terrestrial ecosystems, has significant impact on atmospheric CO2 concentration and climatic dynamics. Nitrogen (N) fertilizer has been heavily applied in agroecosystems at the global scale for high crop yields, and plays a major role in regulating RS. Although the respective response of soil biochemical property and RS to N addition has been widely studied, the contributions of soil biochemical parameters especially in the rhizosphere to changes in RS and its components (soil heterotrophic (RH) and autotrophic (RA) respiration) under N application remain poorly understood. The present study aimed to examine whether the rhizosphere effect alters the relationship between soil biochemical properties and RS under N addition in croplands.
Liang J, Xia J, Shi Z, Jiang L, Ma S, Lu X, Mauritz M, Natali SM, Pegoraro E, Penton CR, Plaza C, Salmon VG, Celis G, Cole JR, Konstantinidis KT, Tiedje JM, Zhou J, Schuur EAG, Luo Y (2018) Biotic responses buffer warming‐induced soil organic carbon loss in Arctic tundra. Global Change Biology 24(10): 4946-4959.
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Read PublicationClimate warming can result in both abiotic (e.g., permafrost thaw) and biotic (e.g., microbial functional genes) changes in Arctic tundra. Recent research has incorporated dynamic permafrost thaw in Earth system models (ESMs) and indicates that Arctic tundra could be a significant future carbon (C) source due to the enhanced decomposition of thawed deep soil C. However, warming‐induced biotic changes may influence biologically related parameters and the consequent projections in ESMs. How model parameters associated with biotic responses will change under warming and to what extent these changes affect projected C budgets have not been carefully examined. In this study, we synthesized six data sets over 5 years from a soil warming experiment at the Eight Mile Lake, Alaska, into the Terrestrial ECOsystem (TECO) model with a probabilistic inversion approach. The TECO model used multiple soil layers to track dynamics of thawed soil under different treatments. Our results show that warming increased light use efficiency of vegetation photosynthesis but decreased baseline (i.e., environment‐corrected) turnover rates of SOC in both the fast and slow pools in comparison with those under control. Moreover, the parameter changes generally amplified over time, suggesting processes of gradual physiological acclimation and functional gene shifts of both plants and microbes. The TECO model predicted that field warming from 2009 to 2013 resulted in cumulative C losses of 224 or 87 g/m<sup>2</sup>, respectively, without or with changes in those parameters. Thus, warming‐induced parameter changes reduced predicted soil C loss by 61%. Our study suggests that it is critical to incorporate biotic changes in ESMs to improve the model performance in predicting C dynamics in permafrost regions.
Liang J, Zhou Z, Huo C, Shi Z, Cole JR, Huang L, Konstantinidis KT, Li X, Liu B, Luo Z, Penton CR, Schuur EAG, Tiedje JM, Wang YP, Wu L, Xia J, Zhou J, Luo Y (2018) More replenishment than priming loss of soil organic carbon with additional carbon input. Nature Communications 9(1):3175.
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Read PublicationIncreases in carbon (C) inputs to soil can replenish soil organic C (SOC) through various mechanisms. However, recent studies have suggested that the increased C input can also stimulate the decomposition of old SOC via priming. Whether the loss of old SOC by priming can override C replenishment has not been rigorously examined. Here we show, through data–model synthesis, that the magnitude of replenishment is greater than that of priming, resulting in a net increase in SOC by a mean of 32% of the added new C. The magnitude of the net increase in SOC is positively correlated with the nitrogen-to-C ratio of the added substrates. Additionally, model evaluation indicates that a two-pool interactive model is a parsimonious model to represent the SOC decomposition with priming and replenishment. Our findings suggest that increasing C input to soils likely promote SOC accumulation despite the enhanced decomposition of old C via priming.
Loranty M, Davydov S, Kropp H, Alexander H, Mack M, Natali S, Zimov N (2018) Vegetation indices do not capture forest cover variation in upland Siberian larch forests. Remote Sensing 10(11).
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Download .PDFBoreal forests are changing in response to climate, with potentially important feedbacks to regional and global climate through altered carbon cycle and albedo dynamics. These feedback processes will be affected by vegetation changes, and feedback strengths will largely rely on the spatial extent and timing of vegetation change. Satellite remote sensing is widely used to monitor vegetation dynamics, and vegetation indices (VIs) are frequently used to characterize spatial and temporal trends in vegetation productivity. In this study we combine field observations of larch forest cover across a 25 km2 upland landscape in northeastern Siberia with high-resolution satellite observations to determine how the Normalized Difference Vegetation Index (NDVI) and the Enhanced Vegetation Index (EVI) are related to forest cover. Across 46 forest stands ranging from 0% to 90% larch canopy cover, we find either no change, or declines in NDVI and EVI derived from PlanetScope CubeSat and Landsat data with increasing forest cover. In conjunction with field observations of NDVI, these results indicate that understory vegetation likely exerts a strong influence on vegetation indices in these ecosystems. This suggests that positive decadal trends in NDVI in Siberian larch forests may correspond primarily to increases in understory productivity, or even to declines in forest cover. Consequently, positive NDVI trends may be associated with declines in terrestrial carbon storage and increases in albedo, rather than increases in carbon storage and decreases in albedo that are commonly assumed. Moreover, it is also likely that important ecological changes such as large changes in forest density or variable forest regrowth after fire are not captured by long-term NDVI trends.
Lu X, Vitousek PM, Mao Q, Gilliam FS, Luo Y, Zhou G, Zou X, Bai E, Scalnon TM, Hou E, Mo J (2018) Plant acclimation to long-term high nitrogen deposition in an N-rich tropical forest. Proceedings of the National Academy of Sciences 115(20):5187.
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Read PublicationElevated atmospheric N deposition threatens ecosystem health through eutrophication in terrestrial ecosystems, but little is known about consequences of N deposition in N-rich tropical ecosystems. We added several levels of N to an N-rich tropical forest and monitored plant growth dynamics, forest nutrient status, plant water use, and water losses from the ecosystem for a decade. We found that plants can acclimate and maintain nutrient balance by altering hydrological cycling. These results demonstrate that while elevated N deposition to already N-rich tropical forests may have minor effects on forest growth, it can exert a detectable influence on hydrological dynamics. Reduced runoff may threaten water supply in rapidly developing tropical regions.Anthropogenic nitrogen (N) deposition has accelerated terrestrial N cycling at regional and global scales, causing nutrient imbalance in many natural and seminatural ecosystems. How added N affects ecosystems where N is already abundant, and how plants acclimate to chronic N deposition in such circumstances, remains poorly understood. Here, we conducted an experiment employing a decade of N additions to examine ecosystem responses and plant acclimation to added N in an N-rich tropical forest. We found that N additions accelerated soil acidification and reduced biologically available cations (especially Ca and Mg) in soils, but plants maintained foliar nutrient supply at least in part by increasing transpiration while decreasing soil water leaching below the rooting zone. We suggest a hypothesis that cation-deficient plants can adjust to elevated N deposition by increasing transpiration and thereby maintaining nutrient balance. This result suggests that long-term elevated N deposition can alter hydrological cycling in N-rich forest ecosystems.
Lu X, Wang YP, Luo Y, Jiang L (2018) Ecosystem carbon transit versus turnover times in response to climate warming and rising atmospheric CO2 concentration. Biogeosciences 15(21): 6559-6572.
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Read PublicationEcosystem carbon (C) transit time is a critical diagnostic parameter to characterize land C sequestration. This parameter has different variants in the literature, including a commonly used turnover time. However, we know little about how different transit time and turnover time are in representing carbon cycling through multiple compartments under a non-steady state. In this study, we estimate both C turnover time as defined by the conventional stock over flux and mean C transit time as defined by the mean age of C mass leaving the system. We incorporate them into the Community Atmosphere Biosphere Land Exchange (CABLE) model to estimate C turnover time and transit time in response to climate warming and rising atmospheric [CO<sub>2</sub>]. Modelling analysis shows that both C turnover time and transit time increase with climate warming but decrease with rising atmospheric [CO<sub>2</sub>]. Warming increases C turnover time by 2.4 years and transit time by 11.8 years in 2100 relative to that at steady state in 1901. During the same period, rising atmospheric [CO<sub>2</sub>] decreases C turnover time by 3.8 years and transit time by 5.5 years. Our analysis shows that 65% of the increase in global mean C transit time with climate warming results from the depletion of fast-turnover C pool. The remaining 35% increase results from accompanied changes in compartment C age structures. Similarly, the decrease in mean C transit time with rising atmospheric [CO<sub>2</sub>] results approximately equally from replenishment of C into fast-turnover C pool and subsequent decrease in compartment C age structure. Greatly different from the transit time, the turnover time, which does not account for changes in either C age structure or composition of respired C, underestimated impacts of warming and rising atmospheric [CO<sub>2</sub>] on C diagnostic time and potentially led to deviations in estimating land C sequestration in multi-compartmental ecosystems.
Luo Y, El-Madeny ST, Filippa G, Ma X, Ahrens B, Carrara A, Gonzalez-Cascon R, Cremonese E, Galvagno M, Hammer TW, Pacheco-Labrador J, Martin PM, Moreno G, Perez-Priego O, Reichstein M, Richardson AD, Romermann C, Migliavacca M (2018) Using near-infrared-enabled digital repeat photography to track structural and physiological phenology in Mediterranean tree-grass ecosystems. Remote Sensing 10(8).
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Download .PDFTree-grass ecosystems are widely distributed. However, their phenology has not yet been fully characterized. The technique of repeated digital photographs for plant phenology monitoring (hereafter referred as PhenoCam) provide opportunities for long-term monitoring of plant phenology, and extracting phenological transition dates (PTDs, e.g., start of the growing season). Here, we aim to evaluate the utility of near-infrared-enabled PhenoCam for monitoring the phenology of structure (i.e., greenness) and physiology (i.e., gross primary productivity-GPP) at four tree-grass Mediterranean sites. We computed four vegetation indexes (VIs) from PhenoCams: (1) green chromatic coordinates (GCC), (2) normalized difference vegetation index (CamNDVI), (3) near-infrared reflectance of vegetation index (CamNIRv), and (4) ratio vegetation index (CamRVI). GPP is derived from eddy covariance flux tower measurement. Then, we extracted PTDs and their uncertainty from different VIs and GPP. The consistency between structural (VIs) and physiological (GPP) phenology was then evaluated. CamNIRv is best at representing the PTDs of GPP during the Green-up period, while CamNDVI is best during the Dry-down period. Moreover, CamNIRv outperforms the other VIs in tracking growing season length of GPP. In summary, the results show it is promising to track structural and physiology phenology of seasonally dry Mediterranean ecosystem using near-infrared-enabled PhenoCam. We suggest using multiple VIs to better represent the variation of GPP.
Mau RL, Dijkstra P, Schwartz E, Koch BJ, Hungate BA (2018) Warming induced changes in soil carbon and nitrogen influence priming responses in four ecosystems. Applied Soil Ecology 124:110-116.
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Read PublicationSoil contains the largest terrestrial pool of carbon (C), but how this pool will be affected by global change remains unknown. Warmer temperatures generally increase soil respiration, while additional C inputs from plants to soil can increase or decrease soil C decomposition rates through a phenomenon known as priming. Priming occurs when soil organic matter (SOM) decomposition rates change in response to a fresh substrate, though the mechanisms underlying priming are poorly understood. Here, we measured priming in four ecosystems during a seven-week incubation with weekly glucose additions. Soil was collected from field warming experiments in the four ecosystems, so our experiment assessed the influence of long-term warming on priming. All treatments exhibited negative priming (reduced SOM decomposition) after the first substrate pulse. Subsequent substrate pulses elicited variable responses, and the effect of long-term warming on priming was ecosystem-dependent. Priming was correlated with changes in soil C and N in response to warming: ecosystems that lost soil C and N over nine years of experimental warming exhibited low rates of priming (decreased SOM decomposition), while ecosystems that gained soil C and N in response to warming had high priming. Consequently, priming may accelerate C losses in ecosystems that exhibit warming-induced C increases, and vice versa, thus partially buffering soil C content against change.
Mauritz M, Celis G, Ebert C, Hutchings J, Ledman J, Natali SM, Pegoraro E, Salmon VG, Schadel C, Taylor M, Schuur EAG (2018) Using stable carbon isotopes of seasonal ecosystem respiration to determine permafrost carbon loss. Biogeosciences.
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Read PublicationHigh latitude warming and permafrost thaw will expose vast stores of deep soil organic carbon (SOC) to decomposition. Thaw also changes water movement causing either wetter or drier soil. The fate of deep SOC under different thaw and moisture conditions is unclear. We measured weekly growing-season δ13C of ecosystem respiration (Recoδ13C) across thaw and moisture conditions (Shallow-Dry; Deep-Dry; Deep-Wet) in a soil warming manipulation. Deep SOC loss was inferred from known δ13C signatures of plant shoot, root, surface soil, and deep soil respiration. In addition, a 2-year-old vegetation removal treatment (No Veg) was used to isolate surface and deep SOC decomposition contributions to Reco. In No Veg, seasonal Recoδ13C indicated that deep SOC loss increased as the soil column thawed, while in vegetated areas, root contributions appeared to dominate Reco. The Recoδ13C differences between Shallow-Dry and Deep-Dry were significant but surprisingly small. This most likely suggests that, under dry conditions, soil warming stimulates root and surface SOC respiration with a negative 13C signature that opposes the more positive 13C signal from increased deep SOC respiration. In Deep-Wet conditions, Recoδ13C suggests reduced deep SOC loss but could also reflect altered diffusion or methane (CH4) dynamics. Together, these results demonstrate that frequent Recoδ13C measurements can detect deep SOC loss and that plants confound the signal. In future studies, soil profile δ13C measurements, vegetation removal across thaw gradients, and isotopic effects of CH4 dynamics could further deconvolute deep SOC loss via surface Reco.
McGuire AD, Lawrence DM, Koven C, Clein JS, Burke E, Chen G, Jafarov E, MacDougall AH, Marchenko S, Nicolsky D, Peng S, Rinke A, Ciais P, Gouttevin I, Hayes DJ, Ji D, Krinner G, Moore JC, Romanovsky V, Schädel C, Schaefer K, Schuur EAG, Zhuang Qianlai (2018) Dependence of the evolution of carbon dynamics in the northern permafrost region on the trajectory of climate change. PNAS 201719903.
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Read PublicationWe conducted a model-based assessment of changes in permafrost area and carbon storage for simulations driven by RCP4.5 and RCP8.5 projections between 2010 and 2299 for the northern permafrost region. All models simulating carbon represented soil with depth, a critical structural feature needed to represent the permafrost carbon–climate feedback, but that is not a universal feature of all climate models. Between 2010 and 2299, simulations indicated losses of permafrost between 3 and 5 million km^2 for the RCP4.5 climate and between 6 and 16 million km^2 for the RCP8.5 climate. For the RCP4.5 projection, cumulative change in soil carbon varied between 66-Pg C (10^15-g carbon) loss to 70-Pg C gain. For the RCP8.5 projection, losses in soil carbon varied between 74 and 652 Pg C (mean loss, 341 Pg C). For the RCP4.5 projection, gains in vegetation carbon were largely responsible for the overall projected net gains in ecosystem carbon by 2299 (8- to 244-Pg C gains). In contrast, for the RCP8.5 projection, gains in vegetation carbon were not great enough to compensate for the losses of carbon projected by four of the five models; changes in ecosystem carbon ranged from a 641-Pg C loss to a 167-Pg C gain (mean, 208-Pg C loss). The models indicate that substantial net losses of ecosystem carbon would not occur until after 2100. This assessment suggests that effective mitigation efforts during the remainder of this century could attenuate the negative consequences of the permafrost carbon–climate feedback.
Melvin AM, Celis G, Johnstone JF, McGuire AD, Genet H, Schuur EAG, Rupp TS, Mack MC (2018) Fuel-reduction management alters plant composition, carbon and nitrogen pools, and soil thaw in Alaskan boreal forest. Ecological Applications 28(1): 149-161.
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Read PublicationIncreasing wildfire activity in Alaska’s boreal forests has led to greater fuel reduction management. Management has been implemented to reduce wildfire spread, but the ecological impacts of these practices are poorly known. We quantified the effects of handthinning and shearblading on above- and belowground stand characteristics, plant species composition, carbon (C) and nitrogen (N) pools, and soil thaw across 19 sites dominated by black spruce (Picea mariana) in interior Alaska treated 2–12 years prior to sampling. The density of deciduous tree seedlings was significantly higher in shearbladed areas compared to unmanaged forest (6.4 vs. 0.1 stems/m2), and unmanaged stands exhibited the highest mean density of conifer seedlings and layers (1.4 stems/m2). Understory plant community composition was most similar between unmanaged and thinned stands. Shearblading resulted in a near complete loss of aboveground tree biomass C pools while thinning approximately halved the C pool size (1.2 kg C/m2 compared to 3.1 kg C/m2 in unmanaged forest). Significantly smaller soil organic layer (SOL) C and N pools were observed in shearbladed stands (3.2 kg C/m2 and 116.8 g N/m2) relative to thinned (6.0 kg C/m2 and 192.2 g N/m2) and unmanaged (5.9 kg C/m2 and 178.7 g N/m2) stands. No difference in C and N pool sizes in the uppermost 10 cm
of mineral soil was observed among stand types. Total C stocks for measured pools was 2.6 kg C/m2 smaller in thinned stands and 5.8 kg C/m2 smaller in shearbladed stands when compared to unmanaged forest. Soil thaw depth averaged 13 cm deeper in thinned areas and 46 cm deeper in shearbladed areas relative to adjacent unmanaged stands, although variability was high across sites. Deeper soil thaw was linked to shallower SOL depth for unmanaged stands and both management types, however for any given SOL depth, thaw tended to be deeper in shearbladed areas compared to unmanaged forest. These findings indicate that fuel-reduction management alters plant community composition, C and N pools, and soil thaw depth, with consequences for ecosystem structure and function beyond those intended for fire management.
Morrissey EM, Mau RL, Schwartz E, Koch BJ, Hayer M, Hungate BA (2018) Taxonomic patterns in the nitrogen assimilation of soil prokaryotes. Enviornmental Microbiology 20(3): 1112-1119.
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Read PublicationNitrogen (N) is frequently a limiting nutrient in soil; its availability can govern ecosystem functions such as primary production and decomposition. Assimilation of N by microorganisms impacts the availability of N in soil. Despite its established ecological significance, the contributions of microbial taxa to N assimilation are unknown. Here we measure N uptake and use by microbial phylotypes and taxonomic groups within a diverse assemblage of soil microbes through quantitative stable isotope probing (qSIP) with 15N. Following incubation with 15NH+4, distinct patterns of 15N assimilation among taxonomic groups were observed. For instance, glucose addition stimulated 15N assimilation in most members of Actinobacteria and Proteobacteria but generally decreased 15N use by Firmicutes and Bacteriodetes. While NH+4 is considered a preferred and universal source of N to prokaryotes, the majority (Greater than 80%) of N assimilation in our soils could be attributed to a handful of active orders. Characterizing N assimilation of taxonomic groups with 15N qSIP may provide a basis for understanding how microbial community composition influences N availability in the environment.
Niu S, Classen AT, Luo Y (2018) Functional traits along a transect. Functional Ecology 32(1): 4-9.
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Read PublicationFunctional traits, which usually develop over evolutionary time‐scales to maximize plant survivorship and functional performances in changing environment, are important indices to explore how ecosystems respond and adapt to a changing environment. Many ecologists have argued that identifying regional to global‐scale patterns in functional traits, at organismal to ecosystem scales, in combination with responses to environmental changes, is necessary to increase our ability to predict how ecosystems will function in the future (Katja & Jeanfrançois, 2009; Reich, Walters, & Ellsworth, 1997; Wright et al., 2017). Global models that have been used to simulate changes in ecosystem function primarily incorporate biogeochemical and ecophysiological processes (Bonan, 2008; Taylor, Stouffer, & Meehl, 2012). These models, however, could not well predict community compositional change (Fisher et al., 2017). Traditionally, plant functional types have been used to represent community composition changes in dynamic global vegetation models (DGVM). That scheme remains poor at predicting ecosystem functions and their responses to climatic change (Sitch et al., 2008). Recently, trait‐based modelling has emerged as one of the most promising approach to simulation of community dynamics under global changes (Markus, Michael, Mahecha, Jens, & Baldocchi, 2014; van Bodegom, Douma, & Verheijen, 2014; Van Bodegom et al., 2012; Violle, Reich, Pacala, Enquist, & Kattge, 2014). To support this trait‐based modelling, it is urgent to compile empirical evidence, develop comprehensive datasets, and reveal the large‐scale patterns and controlling factors of functional traits and their variations along environmental gradients.
Ochoa-Hueso R, Eldridge DJ, Baquerizo, Soliveres S, Bowker MA, Gross N, Le Bagousse-Pinguet Y, Quero JL, Garcia-Gomez M, Valencia E, Arredondo T, Beinticinco L, Bran D, Cea A, Coaguila D, Dougill AJ, Espinosa CI, Gaitan J, Guuroh RT, Guzman E, Gutierrez JR, Hernandez RM, Huber-Sannwald E, Jeffries T, Linstadter A, Mau RL, Monerris J, Prina A, Pucheta E, Stavi I, Thomas AD, Zaady E, Singh BK, Maestre FT (2018) Soil fungal abundance and plant functional traits drive fertile island formation in global drylands. Journal of Ecology 106(1): 242-253.
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Read Publication1. Dryland vegetation is characterized by discrete plant patches that accumulate and capture soil resources under their canopies. These “fertile islands” are major drivers of dryland ecosystem structure and functioning, yet we lack an integrated understanding of the factors controlling their magnitude and variability at the global scale.
2. We conducted a standardized field survey across 236 drylands from five continents. At each site, we measured the composition, diversity and cover of perennial plants. Fertile island effects were estimated at each site by comparing composite soil samples obtained under the canopy of the dominant plants and in open areas devoid of perennial vegetation. For each sample, we measured 15 soil variables (functions) associated with carbon, nitrogen and phosphorus cycling and used the relative interaction index to quantify the magnitude of the fertile island effect for each function. In 80 sites, we also measured fungal and bacterial abundance (quantitative PCR) and diversity (Illumina MiSeq).
3. The most fertile islands, i.e. those where a higher number of functions were simultaneously enhanced, were found at lower elevation sites with greater soil pH values and sand content under semiarid climates, particularly at locations where the presence of tall woody species with a low‐specific leaf area increased fungal abundance beneath plant canopies, the main direct biotic controller of the fertile island effect in the drylands studied. Positive effects of fungal abundance were particularly associated with greater nutrient contents and microbial activity (soil extracellular enzymes) under plant canopies.
4. Synthesis. Our results show that the formation of fertile islands in global drylands largely depends on: (1) local climatic, topographic and edaphic characteristics, (2) the structure and traits of local plant communities and (3) soil microbial communities. Our study also has broad implications for the management and restoration of dryland ecosystems worldwide, where woody plants are commonly used as nurse plants to enhance the establishment and survival of beneficiary species. Finally, our results suggest that forecasted increases in aridity may enhance the formation of fertile islands in drylands worldwide.
Papp K, Hungate BA, Schwartz E (2018) Microbial rRNA synthesis and growth compared through quantitative stable isotope probing with H218O. Applied and Environmental Microbiology 84(8).
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Read PublicationGrowing bacteria have a high concentration of ribosomes to ensure sufficient protein synthesis, which is necessary for genome replication and cellular division. To elucidate whether metabolic activity of soil microorganisms is coupled with growth, we investigated the relationship between rRNA and DNA synthesis in a soil bacterial community using quantitative stable isotope probing (qSIP) with H2 18O. Most soil bacterial taxa were metabolically active and grew, and there was no significant difference between the isotopic composition of DNA and RNA extracted from soil incubated with H2 18O. The positive correlation between 18O content of DNA and rRNA of taxa, with a slope statistically indistinguishable from 1 (slope = 0.96; 95% confidence interval [CI], 0.90 to 1.02), indicated that few taxa made new rRNA without synthesizing new DNA. There was no correlation between rRNA-to-DNA ratios obtained from sequencing libraries and the atom percent excess (APE) 18O values of DNA or rRNA, suggesting that the ratio of rRNA to DNA is a poor indicator of microbial growth or rRNA synthesis. Our results support the notion that metabolic activity is strongly coupled to cellular division and suggest that nondividing taxa do not dominate soil metabolic activity. IMPORTANCE Using quantitative stable isotope probing of microbial RNA and DNA with H2 18O, we show that most soil taxa are metabolically active and grow because their nucleic acids are significantly labeled with 18O. A majority of the populations that make new rRNA also grow, which argues against the common paradigm that most soil taxa are dormant. Additionally, our results indicate that relative sequence abundance-based RNA-to-DNA ratios, which are frequently used for identifying active microbial populations in the environment, underestimate the number of metabolically active taxa within soil microbial communities.
Papp K, Mau RL, Hayer M, Koch BJ, Hungate BA, Schwartz E (2018) Quantitative stable isotope probing with H218O reveals that most bacterial taxa in soil synthesize new ribosomal RNA. The ISME Journal.
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Read PublicationMost soil bacterial taxa are thought to be dormant, or inactive, yet the extent to which they synthetize new rRNA is poorly understood. We analyzed 18O composition of RNA extracted from soil incubated with H218O and used quantitative stable isotope probing to characterize rRNA synthesis among microbial taxa. RNA was not fully labeled with 18O, peaking at a mean of 23.6 ± 6.8 atom percent excess (APE) 18O after eight days of incubation, suggesting some ribonucleotides in soil were more than eight days old. Microbial taxa varied in the degree they incorporated 18O into their rRNA over time and there was no correlation between the APE 18O of bacterial rRNA and their rRNA to DNA ratios, suggesting that the ratios were not appropriate to measure ribonucleotide synthesis. Our study indicates that, on average, 94% of soil taxa produced new rRNA and therefore were metabolically active.
Pegoraro E, Mauritz M, Bracho R, Ebert C, Dijkstra P, Hungate BA, Konstantinidis KT, Luo Y, Schädel C, Tiedje JM, Zhou J, Schuur EAG (2018) Glucose addition increases the magnitude and decreases the age of soil respired carbon in a long-term permafrost incubation study. Soil Biology and Biochemistry.
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Read PublicationHigher temperatures in northern latitudes will increase permafrost thaw and stimulate above-and belowground plant biomass growth in tundra ecosystems. Higher plant productivity increases the input of easily decomposable carbon (C) to soil, which can stimulate microbial activity and increase soil organic matter decomposition rates. This phenomenon, known as the priming effect, is particularly interesting in permafrost because an increase in C supply to deep, previously frozen soil may accelerate decomposition of C stored for hundreds to thousands of years. The sensitivity of old permafrost C to priming is not well known; most incubation studies last less than one year, and so focus on fast-cycling C pools. Furthermore, the age of respired soil C is rarely measured, even though old C may be vulnerable to labile C inputs. We incubated soil from a moist acidic tundra site in Eight Mile Lake, Alaska for 409 days at 15 °C. Soil from surface (0–25 cm), transition (45–55 cm), and permafrost (65–85 cm) layers were amended with three pulses of uniformly 13C labeled glucose or cellulose, every 152 days. Glucose addition resulted in positive priming in the permafrost layer 7 days after each substrate addition, eliciting a two-fold increase in cumulative soil C loss relative to unamended soils with consistent effects across all three pulses. In the transition and permafrost layers, glucose addition significantly decreased the age of soil-respired CO2C with Δ14C values that were 115‰ higher. Previous field studies that measured the age of respired C in permafrost regions have attributed younger Δ14C ecosystem respiration values to higher plant contributions. However, the results from this study suggest that positive priming, due to an increase in fresh C supply to deeply thawed soil layers, can also explain the respiration of younger C observed at the ecosystem scale. We must consider priming effects to fully understand permafrost C dynamics, or we risk underestimating the contribution of soil C to ecosystem respiration.
Plaza C, Zaccone C, Sawicka K, Mendez AM, Tarquis A, Gasco G, Heuvelink GBM, Schuur EAG, Maestre FT (2018) Soil resources and element stocks in drylands to face global issues. Scientific Reports 8(1):13788.
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Read PublicationDrylands (hyperarid, arid, semiarid, and dry subhumid ecosystems) cover almost half of Earth’s land surface and are highly vulnerable to environmental pressures. Here we provide an inventory of soil properties including carbon (C), nitrogen (N), and phosphorus (P) stocks within the current boundaries of drylands, aimed at serving as a benchmark in the face of future challenges including increased population, food security, desertification, and climate change. Aridity limits plant production and results in poorly developed soils, with coarse texture, low C:N and C:P, scarce organic matter, and high vulnerability to erosion. Dryland soils store 646 Pg of organic C to 2 m, the equivalent of 32% of the global soil organic C pool. The magnitude of the historic loss of C from dryland soils due to human land use and cover change and their typically low C:N and C:P suggest high potential to build up soil organic matter, but coarse soil textures may limit protection and stabilization processes. Restoring, preserving, and increasing soil organic matter in drylands may help slow down rising levels of atmospheric carbon dioxide by sequestering C, and is strongly needed to enhance food security and reduce the risk of land degradation and desertification.
Rasmussen C, Heckman K, Wieder WR, Keiluweit M, Lawrence CR, Berhe AA, Blankinship JC, Crow SE, Druhan JL, Hicks Pries CE, Marin-Spiotta E, Plante AF, Schädel C, Schimel JP, Sierra CA, Thompson A, Wagai R (2018) Beyond clay: Towards an improved set of variables for predicting soil organic matter content. Biogeochemistry 137(3): 297-306.
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Read PublicationImproved quantification of the factors controlling soil organic matter (SOM) stabilization at continental to global scales is needed to inform projections of the largest actively cycling terrestrial carbon pool on Earth, and its response to environmental change. Biogeochemical models rely almost exclusively on clay content to modify rates of SOM turnover and fluxes of climate-active CO2 to the atmosphere. Emerging conceptual understanding, however, suggests other soil physicochemical properties may predict SOM stabilization better than clay content. We addressed this discrepancy by synthesizing data from over 5,500 soil profiles spanning continental scale environmental gradients. Here, we demonstrate that other physicochemical parameters are much stronger predictors of SOM content, with clay content having relatively little explanatory power. We show that exchangeable calcium strongly predicted SOM content in water-limited, alkaline soils, whereas with increasing moisture availability and acidity, iron- and aluminum-oxyhydroxides emerged as better predictors, demonstrating that the relative importance of SOM stabilization mechanisms scales with climate and acidity. These results highlight the urgent need to modify biogeochemical models to better reflect the role of soil physicochemical properties in SOM cycling.
Richardson AD (2018) Tracking seasonal rhythms of plants in diverse ecosystems with digital camera imagery. New Phytologist.
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Read PublicationGlobal change is shifting the seasonality of vegetation in ecosystems around the globe. High-frequency digital camera imagery, and vegetation indices derived from that imagery, is facilitating better tracking of phenological responses to environmental variation. This method, commonly referred to as the “phenocam” approach, is well-suited to several specific applications, including: close-up observation of individual organisms; long-term canopy-level monitoring at individual sites; automated regional-to-continental scale observatory networks; and tracking responses to experimental treatments. Several camera networks are already well-established, and some camera records are a more than a decade long. These data can be used to identify the environmental controls on phenology in different ecosystems, which will contribute to the development of improved prognostic phenology models. This article is protected by copyright. All rights reserved.
Richardson AD, Hufkens K, Milliman T, Aubrecht DM, Chen M, Gray JM, Johnston MR, Keenan TF, Klosterman ST, Kosmala M, Melaas EK, Friedl MA, Frolking S (2018) Tracking vegetation phenology across diverse North American biomes using PhenoCam imagery. Nature 5:180028.
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Read PublicationVegetation phenology controls the seasonality of many ecosystem processes, as well as numerous biosphere-atmosphere feedbacks. Phenology is also highly sensitive to climate change and variability. Here we present a series of datasets, together consisting of almost 750 years of observations, characterizing vegetation phenology in diverse ecosystems across North America. Our data are derived from conventional, visible-wavelength, automated digital camera imagery collected through the PhenoCam network. For each archived image, we extracted RGB (red, green, blue) colour channel information, with means and other statistics calculated across a region-of-interest (ROI) delineating a specific vegetation type. From the high-frequency (typically, 30 min) imagery, we derived time series characterizing vegetation colour, including “canopy greenness”, processed to 1- and 3-day intervals. For ecosystems with one or more annual cycles of vegetation activity, we provide estimates, with uncertainties, for the start of the “greenness rising” and end of the “greenness falling” stages. The database can be used for phenological model validation and development, evaluation of satellite remote sensing data products, benchmarking earth system models, and studies of climate change impacts on terrestrial ecosystems.
Richardson AD, Hufkens K, Milliman T, Aubrecht DM, Furze ME, Seyednasrollah B, Krassovski MB, Latimer JM, Nettles WR, Heiderman RR, Warren JM, Hanson PJ (2018) Ecosystem warming extends vegetation activity but heightens vulnerability to cold temperatures. Nature.
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Read PublicationShifts in vegetation phenology are a key example of the biological effects of climate change. However, there is substantial uncertainty about whether these temperature-driven trends will continue, or whether other factors—for example, photoperiod—will become more important as warming exceeds the bounds of historical variability. Here we use phenological transition dates derived from digital repeat photography to show that experimental whole-ecosystem warming treatments of up to +9 °C linearly correlate with a delayed autumn green-down and advanced spring green-up of the dominant woody species in a boreal <i>Picea</i>–<i>Sphagnum</i> bog. Results were confirmed by direct observation of both vegetative and reproductive phenology of these and other bog plant species, and by multiple years of observations. There was little evidence that the observed responses were constrained by photoperiod. Our results indicate a likely extension of the period of vegetation activity by 1–2 weeks under a ‘CO<sub>2</sub> stabilization’ climate scenario (+2.6 ± 0.7 °C), and 3–6 weeks under a ‘high-CO<sub>2</sub> emission’ scenario (+5.9 ± 1.1 °C), by the end of the twenty-first century. We also observed severe tissue mortality in the warmest enclosures after a severe spring frost event. Failure to cue to photoperiod resulted in precocious green-up and a premature loss of frost hardiness, which suggests that vulnerability to spring frost damage will increase in a warmer world. Vegetation strategies that have evolved to balance tradeoffs associated with phenological temperature tracking may be optimal under historical climates, but these strategies may not be optimized for future climate regimes. These in situ experimental results are of particular importance because boreal forests have both a circumpolar distribution and a key role in the global carbon cycle.
Richardson AD, Hufkens K, Milliman T, Frolking S (2018) Intercomparison of phenological transition dates derived from the PhenoCam Dataset V1.0 and MODIS satellite remote sensing. Scientific Reports 8: 5679.
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Read PublicationPhenology is a valuable diagnostic of ecosystem health, and has applications to environmental monitoring and management. Here, we conduct an intercomparison analysis using phenological transition dates derived from near-surface PhenoCam imagery and MODIS satellite remote sensing. We used approximately 600 site-years of data, from 128 camera sites covering a wide range of vegetation types and climate zones. During both “greenness rising” and “greenness falling” transition phases, we found generally good agreement between PhenoCam and MODIS transition dates for agricultural, deciduous forest, and grassland sites, provided that the vegetation in the camera field of view was representative of the broader landscape. The correlation between PhenoCam and MODIS transition dates was poor for evergreen forest sites. We discuss potential reasons (including sub-pixel spatial heterogeneity, flexibility of the transition date extraction method, vegetation index sensitivity in evergreen systems, and PhenoCam geolocation uncertainty) for varying agreement between time series of vegetation indices derived from PhenoCam and MODIS imagery. This analysis increases our confidence in the ability of satellite remote sensing to accurately characterize seasonal dynamics in a range of ecosystems, and provides a basis for interpreting those dynamics in the context of tangible phenological changes occurring on the ground.
Rubin R, Roybal CM (2018) Plant community responses to mastication and mulching of one-seed juniper (Juniperus monosperma). Rangeland Ecology & Management 71(6): 753-756.
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Read PublicationMechanical cutting and mastication of juniper trees aims to restore grassland habitat by reducing the density of encroaching woody species. However, the associated soil disturbance may also create conduits for invasive species, a risk that must be mitigated by land managers. We characterized herbaceous communities in treated and adjacent untreated areas in a piñon-juniper (Pinus edulis and Juniper monosperma) woodland in northern Arizona 2.5 years after treatment. Untreated plots had 4× the herbaceous cover (82%) than treated plots (21%). Within treated plots, native species cover (19%) was 10× higher than invasive species cover (2%). Furthermore, treated plots exhibited greater plant community variability and diversity than untreated plots, driven by an increase in the diversity of native grasses and non-native forbs. No new recruits were Arizona listed noxious weeds, indicating that, at least in the short term, mastication is not producing invasive species hot spots in this piñon-juniper woodland.
Rubin RL, Koch GW, Martinez A, Mau RL, Bowker MA, Hungate BA (2018) Developing climate-smart restoration: Can plant microbiomes be hardened against heat waves?. Ecological Applications 28(6):1594-1605.
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Read PublicationHeat waves are increasing in frequency and intensity, presenting a challenge for the already difficult practice of ecological restoration. We investigated whether pre-heating locally sourced rhizosphere soil (inoculum) could acclimatize plants to a field-imposed heat wave in a restoration setting. Soil heating in the laboratory caused a marked shift in rhizosphere bacterial community composition, accompanied by an increase in species evenness. Furthermore, pre-heated rhizosphere soil reduced plant height, number of leaves, and shoot mass of the C4 grass, blue grama (Bouteloua gracilis), and it reduced the shoot mass of the C3 grass, Arizona fescue (Festuca arizonica) in the glasshouse. Following transplantation and the application of a field heat wave, pre-heated inoculum did not influence heat wave survival for either plant species. However, there were strong species-level responses to the field heat wave. For instance, heat wave survivorship was over four times higher in blue grama (92%) than in Arizona fescue (22%). These results suggest that the use of C4 seeds may be preferable for sites exhibiting high heat wave risk. Further research is needed to understand whether inocula are more effective in highly degraded soil in comparison with partially degraded soils.
Ryan EM, Ogle K, Kropp H, Samuels-Crow KE, Carrillo Y, Pendall E (2018) Modeling soil CO2 production and transport with dynamic source and diffusion terms: Testing the steady-state assumption using DETECT v1.0. Geoscientific Model Development 11(5): 1909-1928.
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Read PublicationThe flux of CO<sub>2</sub> from the soil to the atmosphere (soil respiration, <i>R</i><sub>soil</sub>) is a major component of the global carbon (C) cycle. Methods to measure and model <i>R</i><sub>soil</sub>, or partition it into different components, often rely on the assumption that soil CO<sub>2</sub> concentrations and fluxes are in steady state, implying that <i>R</i><sub>soil</sub> is equal to the rate at which CO<sub>2</sub> is produced by soil microbial and root respiration. Recent research, however, questions the validity of this assumption. Thus, the aim of this work was two-fold: (1) to describe a non-steady state (NSS) soil CO<sub>2</sub> transport and production model, DETECT, and (2) to use this model to evaluate the environmental conditions under which <i>R</i><sub>soil</sub>and CO<sub>2</sub> production are likely in NSS. The backbone of DETECT is a non-homogeneous, partial differential equation (PDE) that describes production and transport of soil CO<sub>2</sub>, which we solve numerically at fine spatial and temporal resolution (e.g., 0.01m increments down to 1m, every 6h). Production of soil CO<sub>2</sub> is simulated for every depth and time increment as the sum of root respiration and microbial decomposition of soil organic matter. Both of these factors can be driven by current and antecedent soil water content and temperature, which can also vary by time and depth. We also analytically solved the ordinary differential equation (ODE) corresponding to the steady-state (SS) solution to the PDE model. We applied the DETECT NSS and SS models to the six-month growing season period representative of a native grassland in Wyoming. Simulation experiments were conducted with both model versions to evaluate factors that could affect departure from SS, such as (1) varying soil texture; (2) shifting the timing or frequency of precipitation; and (3) with and without the environmental antecedent drivers. For a coarse-textured soil, <i>R</i><sub>soil</sub> from the SS model closely matched that of the NSS model. However, in a fine-textured (clay) soil, growing season <i>R</i><sub>soil</sub> was ∼ 3% higher under the assumption of NSS (versus SS). These differences were exaggerated in clay soil at daily time scales whereby <i>R</i><sub>soil</sub> under the SS assumption deviated from NSS by up to 35% on average in the 10 days following a major precipitation event. Incorporation of antecedent drivers increased the magnitude of <i>R</i><sub>soil</sub> by 15 to 37% for coarse- and fine-textured soils, respectively. However, the responses of <i>R</i><sub>soil</sub> to the timing of precipitation and antecedent drivers did not differ between SS and NSS assumptions. In summary, the assumption of SS conditions can be violated depending on soil type and soil moisture status, as affected by precipitation inputs. The DETECT model provides a framework for accommodating NSS conditions to better predict <i>R</i><sub>soil</sub> and associated soil carbon cycling processes.
Sabo JL, Caron M, Doucett R, Dibble KL, Ruhi A, Marks JC, Hungate BA, Kennedy TA (2018) Pulsed flows, tributary inputs and food‐web structure in a highly regulated river. Journal of Applied Ecology.
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Read Publication1.Dams disrupt the river continuum, altering hydrology, biodiversity and energy flow. Although research indicates that tributary inputs have the potential to dilute these effects, knowledge at the food‐web level is still scarce.
2.Here, we examined the riverine food‐web structure of the Colorado River below Glen Canyon Dam, focusing on organic matter sources, trophic diversity and food chain length. We asked how these components respond to pulsed flows from tributaries following monsoon thunderstorms that seasonally increase streamflow in the American Southwest.
3.Tributaries increased the relative importance of terrestrial organic matter, particularly during the wet season below junctures of key tributaries. This contrasted with the algal‐based food‐web present immediately below Glen Canyon Dam.
4.Tributary inputs during the monsoon also increased trophic diversity and food chain length: food chain length peaked below the confluence with the largest tributary (by discharge) in Grand Canyon, increasing by greater than 1 trophic level over a 4–5 km reach possibly due to aquatic prey being flushed into the mainstem during heavy rain events.
5.Our results illustrate that large tributaries can create seasonal discontinuities, influencing riverine food‐web structure in terms of allochthony, food‐web diversity and food chain length.
6.Synthesis and applications. Pulsed flows from unregulated tributaries following seasonal monsoon rains increase the importance of terrestrially derived organic matter in large, regulated river food webs, increasing food chain length and trophic diversity downstream of tributary inputs. Protecting unregulated tributaries within hydropower cascades may be important if we are to mitigate food‐web structure alteration due to flow regulation by large dams. This is critical in the light of global hydropower development, especially in megadiverse, developing countries where dam placement (including completed and planned structures) is in tributaries.
Salmon VG, Schädel C, Bracho R, Pegoraro E, Celis G, Mauritz M, Mack MC, Schuur EAG (2018) Adding depth to our understanding of nitrogen dynamics in permafrost soils. Journal of Geophysical Research: Biogeosciences 123(8):2497-2512.
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Read PublicationLosses of C from decomposing permafrost may be offset by increased productivity of tundra plants, but nitrogen availability partially limits plant growth in tundra ecosystems. In this soil incubation experiment carbon (C) and nitrogen (N) cycling dynamics were examined from the soil surface down through upper permafrost. We found that losses of CO2 were negatively correlated to net N mineralization because C-rich surface soils mineralized little N, while deep soils had low rates of C respiration but high rates of net N mineralization. Permafrost soils released a large flush of inorganic N when initially thawed. Depth-specific rates of N mineralization from the incubation were combined with thaw depths and soil temperatures from a nearby manipulative warming experiment to simulate the potential magnitude, timing, and depth of inorganic N release during the process of permafrost thaw. Our calculations show that inorganic N released from newly thawed permafrost may be similar in magnitude to the increase in N mineralized by warmed soils in the middle of the profile. The total release of inorganic N from the soil profile during the simulated thaw process was twice the size of the observed increase in the foliar N pool observed at the manipulative experiment. Our findings suggest that increases in N availability are likely to outpace the N demand of tundra plants during the first 5 years of permafrost thaw and may increase C losses from surface soils as well as induce denitrification and leaching of N from these ecosystems.
Samuels-Crow KE, Ryan E, Pendall E, Ogle K (2018) Temporal coupling of subsurface and surface soil CO2 fluxes: Insights from a nonsteady state model and cross-wavelet coherence analysis. Journal of Geophysical Research: Biogeosciences 123(4): 1406-1424.
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Read PublicationInferences about subsurface CO2 fluxes often rely on surface soil respiration (Rsoil) estimates because directly measuring subsurface microbial and root respiration (collectively, CO2 production, STotal) is difficult. To evaluate how well Rsoil serves as a proxy for STotal, we applied the nonsteady state DEconvolution of Temporally varying Ecosystem Carbon componenTs model (0.01-m vertical resolution), using 6-hourly data from a Wyoming grassland, in six simulations that cross three soil types (clay, sandy loam, and sandy) with two depth distributions of subsurface biota. We used cross-wavelet coherence analysis to examine temporal coherence (localized linear correlation) and offsets (lags) between STotal and Rsoil and fluxes and drivers (e.g., soil temperature and moisture). Cross-wavelet coherence revealed higher coherence between fluxes and drivers than linear regressions between concurrent variables. Soil texture and moisture exerted the strongest controls over coherence between CO2 fluxes. Coherence between CO2 fluxes in all soil types was strong at short (~1 day) and long periods (>8 days), but soil type controlled lags, and rainfall events decoupled the fluxes at periods of 1?8 days for several days in sandy soil, up to 1 week in sandy loam, and for a month or more in clay soil. Concentrating root and microbial biomass nearer the surface decreased lags in all soil types and increased coherence up to 10% in clay soil. The assumption of high temporal coherence between Rsoil and STotal is likely valid in dry, sandy soil, but may lead to underestimates of short-term STotal in semiarid grasslands with fine-grained and/or wet soil.
Schädel C, Koven CD, Lawrence DM, Celis G, Garnello AJ, Hutchings J, Mauritz M, Natali SM, Pegoraro E, Rodenhizer H, Salmon VG, Taylor MA, Webb EE, Wieder WR, Schuur EAG (2018) Divergent patterns of experimental and model-derived permafrost ecosystem carbon dynamics in response to Arctic warming. Environmental Research Letters 13(10): 105002.
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Read PublicationIn the last few decades, temperatures in the Arctic have increased twice as much as the rest of the globe. As permafrost thaws in response to this warming, large amounts of soil organic matter may become vulnerable to decomposition. Microbial decomposition will release carbon (C) from permafrost soils, however, warmer conditions could also lead to enhanced plant growth and C uptake. Field and modeling studies show high uncertainty in soil and plant responses to climate change but there have been few studies that reconcile field and model data to understand differences and reduce uncertainty. Here, we evaluate gross primary productivity (GPP), ecosystem respiration (R eco ), and net ecosystem C exchange (NEE) from eight years of experimental soil warming in moist acidic tundra against equivalent fluxes from the Community Land Model during simulations parameterized to reflect the field conditions associated with this manipulative field experiment. Over the eight-year experimental period, soil temperatures and thaw depths increased with warming in field observations and model simulations. However, the field and model results do not agree on warming effects on water table depth; warming created wetter soils in the field and drier soils in the models. In the field, initial increases in growing season GPP, R eco , and NEE to experimentally-induced permafrost thaw created a higher C sink capacity in the first years followed by a stronger C source in years six through eight. In contrast, both models predicted linear increases in GPP, R eco , and NEE with warming. The divergence of model results from field experiments reveals the role subsidence, hydrology, and nutrient cycling play in influencing the C flux responses to permafrost thaw, a complexity that the models are not structurally able to predict, and highlight challenges associated with projecting C cycle dynamics across the Arctic.
Schafer JL, Mack MC (2018) Nutrient limitation of plant productivity in scrubby flatwoods: Does fire shift nitrogen versus phosphorus limitation?. Plant Ecology 219(9):1063-1079.
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Read PublicationDifferences in the biogeochemistry of nitrogen (N) and phosphorus (P) lead to differential losses and inputs during and over time after fire such that fire may affect nutrient limitation of primary productivity. We conducted a nutrient addition experiment in scrubby flatwoods, a Florida scrub community type, to test the hypothesis that nutrient limitation of primary productivity shifts from N limitation in recently burned sites to P limitation in longer unburned sites. We added three levels of N, P, and N and P together to sites 6 weeks, 8 years, and 20 years postfire and assessed the effects of nutrient addition on above- and belowground productivity and nutrient concentrations. At the community level, nutrient addition did not affect aboveground biomass, but root productivity increased with high N + P addition in sites 8 and 20 years after fire. At the species level, N addition increased leaf biomass of saw palmetto (Serenoa repens) in sites 6 weeks and 20 years postfire, while P addition increased foliar %P and apical shoot growth of scrub oak (Quercus inopina) in sites 8 and 20 years postfire, respectively. Contrary to our hypothesis, nutrient limitation does not appear to shift with time after fire; recently burned sites show little evidence of nutrient limitation, while increased belowground productivity indicates that scrubby flatwoods are co-limited by N and P at intermediate and longer times after fire.
Schuur EAG & Mack MC (2018) Ecological response to permafrost thaw and consequences for local and global ecosystem services. Annual Review of Ecology, Evolution, and Systematics 49: 279-301.
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Read PublicationThe Arctic may seem remote, but the unprecedented environmental changes occurring there have important consequences for global society. Of all Arctic system components, changes in permafrost (perennially frozen ground) are one of the least documented. Permafrost is degrading as a result of climate warming, and evidence is mounting that changing permafrost will have significant impacts within and outside the region. This review asks: What are key structural and functional properties of ecosystems that interact with changing permafrost, and how do these ecosystem changes affect local and global society? Here, we look beyond the classic definition of permafrost to include a broadened focus on the composition of frozen ground, including the ice and the soil organic carbon content, and how it is changing. This ecological perspective of permafrost serves to identify areas of both vulnerability and resilience as climate, ecological disturbance regimes, and the human footprint all continue to change in this sensitive and critical region of Earth.
Shi Z, Crowell S, Luo Y, Moore B (2018) Model structures amplify uncertainty in predicted soil carbon responses to climate change. Nature Communications 9(1):2171.
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Read PublicationLarge model uncertainty in projected future soil carbon (C) dynamics has been well documented. However, our understanding of the sources of this uncertainty is limited. Here we quantify the uncertainties arising from model parameters, structures and their interactions, and how those uncertainties propagate through different models to projections of future soil carbon stocks. Both the vertically resolved model and the microbial explicit model project much greater uncertainties to climate change than the conventional soil C model, with both positive and negative C-climate feedbacks, whereas the conventional model consistently predicts positive soil C-climate feedback. Our findings suggest that diverse model structures are necessary to increase confidence in soil C projection. However, the larger uncertainty in the complex models also suggests that we need to strike a balance between model complexity and the need to include diverse model structures in order to forecast soil C dynamics with high confidence and low uncertainty.
Shi Z, Lin Y, Wilcox KR, Souza L, Jiang L, Jung CG, Xu X, Yuan M, Guo X, Wu L, Zhou J, Luo Y (2018) Successional change in species composition alters climate sensitivity of grassland productivity. Global Change Biology 24(10):4993-5003.
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Read PublicationSuccession theory predicts altered sensitivity of ecosystem functions to disturbance (i.e., climate change) due to the temporal shift in plant community composition. However, empirical evidence in global change experiments is lacking to support this prediction. Here, we present findings from an 8-year long-term global change experiment with warming and altered precipitation manipulation (double and halved amount). First, we observed a temporal shift in species composition over 8 years, resulting in a transition from an annual C3-dominant plant community to a perennial C4-dominant plant community. This successional transition was independent of any experimental treatments. During the successional transition, the response of aboveground net primary productivity (ANPP) to precipitation addition magnified from neutral to +45.3%, while the response to halved precipitation attenuated substantially from ?17.6% to neutral. However, warming did not affect ANPP in either state. The findings further reveal that the time-dependent climate sensitivity may be regulated by successional change in species composition, highlighting the importance of vegetation dynamics in regulating the response of ecosystem productivity to precipitation change.
Shiga YP, Michalak AM, Yuanyuan F, Schaefer K, Andrews AE, Huntzinger DH, Schwalm CR, Thoning K, Wei Y (2018) Forests dominate the interannual variability of the North American carbon sink. Environmental Research Letters 13(8): 084015.
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Read PublicationUnderstanding what drives the interannual variability (IAV) of the land carbon sink is crucial for improving future predictions of this important, yet uncertain, component of the climate system. While drivers of global and hemispheric-scale net ecosystem exchange (NEE) IAV have been investigated, our understanding of the drivers of NEE IAV at regional scales (e.g. sub-continental, biome-level) is quite poor. Here we explore the biome-level attribution and drivers of North American NEE using inverse estimates derived from a dense network of atmospheric CO 2 observations. We find that deciduous broadleaf and mixed forests are the primary regions responsible for North American NEE IAV, which differs from the ecoregions identified for the globe and Northern Hemisphere. We also find that a suite of terrestrial biosphere models (TBMs) do not agree on the dominant biome contributing to NEE IAV, with TBMs falling along an apparent spectrum ranging between those with IAV dominated primarily by forested ecosystems to those with IAV dominated by non-forested ecosystems. Furthermore, this regional trade-off in TBM NEE IAV is found to be linked to differing regional responses to environmental drivers among TBMs. This work displays the importance of extra-tropical forests in driving continental NEE IAV and also highlights the challenges and limitations of using TBMs to inform regional-scale carbon flux dynamics.
Siders AC, Compson ZG, Hungate BA, Dijkstra P, Koch GW, Wymore AS, Grandy AS, Marks JC (2018) Litter identity affects assimilation of carbon and nitrogen by a shredding caddisfly. Ecosphere 9(7):e02340.
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Read PublicationEcologists often equate litter quality with decomposition rate. In soil and sediments, litter that is rapidly decomposed by microbes often has low concentrations of tannin and lignin and low C:N ratios. Do these same traits also favor element transfer to higher trophic levels in streams, where many insects depend on litter as their primary food source? We test the hypothesis that slow decomposition rates promote element transfer from litter to insects, whereas rapid decomposition favors microbes. We measured carbon and nitrogen fluxes from four plant species to a leaf-shredding caddisfly using isotopically labeled litter. Caddisflies assimilated a higher percentage of litter carbon and nitrogen lost from slowly decomposing litters (Platanus wrightii and Quercus gambelii). In contrast, rapidly decomposing litters (Fraxinus velutina and Populus fremontii) supported higher microbial biomass. These results challenge the view that rapidly decomposing litter is higher quality by demonstrating that slowly decomposing litters provide a critical resource for insects.
Stephens JJ, Black TA, Jassal RS, Nesic Z, Grant NJ, Barr AG, Helgason WD, Richardson AD, Johnson MS, Christen A (2018) Effects of forest tent caterpillar defoliation on carbon and water fluxes in a boreal aspen stand. Agricultural and Forest Meteorology 253: 176-189.
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Read PublicationInsect outbreaks can significantly influence carbon (C) and water balances of forests. Forest tent caterpillars (FTC) (Malacosoma disstria Hübner) are one of the most prominent insects found in aspen forests in Canada and have the potential to considerably influence regional C and water fluxes. In the summer of 2016, an FTC infestation occurred in a ca. 100 -year-old trembling aspen (Populus tremuloides) stand in the southern boreal forest where the longterm research site known as Old Aspen (OA) is located. The infestation led to nearly complete defoliation of the canopy during the leafing out period when photosynthesis, and thus C uptake, is progressing towards maximum levels. We used 21 years of eddy-covariance (EC) and climate measurements covering pre-infestation and infestation periods to estimate the impact of the FTC infestation on net ecosystem production (NEP), gross ecosystem production (GEP) and evapotranspiration (E). Defoliation in 2016 reduced annual NEP to −130 g C m−2 y−1 and GEP to 798 g C m−2 y−1, respectively, which were much less than their 20-year means (NEP = 118 ± 53 g C m−2 y−1, GEP = 1057 ± 74 g C m−2 y−1), and resulted in the most negative annual NEP value of the 21 years of measurements at the OA site. NEP for 2016 was even lower than values observed during three drought years (2001–2003). However, FTC infestation caused little effect on annual E. FTC infestation reduced the near-surface remotely-measured greenness index, green chromatic coordinate (GCC), to ∼0.32 on June 10 in comparison to ∼0.40 in other years. The defoliation, observable from space as reductions in normalized difference vegetation index (NDVI) values, also showed a negligible effect on E but a large effect on the C fluxes.
Sulman BN, Moore JAM, Abramoff R, Averill C, Kivlin S, Georgiou K, Sridhar B, Harman MD, Wang G, Wieder WR, Bradford MA, Luo Y, Mayes MA, Morrison E, Riley WJ, Salazar A, Schimel JP, Tan J, Classen AT (2018) Multiple models and experiments underscore large uncertainty in soil carbon dynamics. Biogeochemistry 141(2): 109-123.
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Read PublicationSoils contain more carbon than plants or the atmosphere, and sensitivities of soil organic carbon (SOC) stocks to changing climate and plant productivity are a major uncertainty in global carbon cycle projections. Despite a consensus that microbial degradation and mineral stabilization processes control SOC cycling, no systematic synthesis of long-term warming and litter addition experiments has been used to test process-based microbe-mineral SOC models. We explored SOC responses to warming and increased carbon inputs using a synthesis of 147 field manipulation experiments and five SOC models with different representations of microbial and mineral processes. Model projections diverged but encompassed a similar range of variability as the experimental results. Experimental measurements were insufficient to eliminate or validate individual model outcomes. While all models projected that CO2 efflux would increase and SOC stocks would decline under warming, nearly one-third of experiments observed decreases in CO2 flux and nearly half of experiments observed increases in SOC stocks under warming. Long-term measurements of C inputs to soil and their changes under warming are needed to reconcile modeled and observed patterns. Measurements separating the responses of mineral-protected and unprotected SOC fractions in manipulation experiments are needed to address key uncertainties in microbial degradation and mineral stabilization mechanisms. Integrating models with experimental design will allow targeting of these uncertainties and help to reconcile divergence among models to produce more confident projections of SOC responses to global changes.
Taylor MA, Celis G, Ledman JD, Bracho R, Schuur EAG (2018) Methane efflux measured by eddy covariance in Alaskan upland tundra undergoing permafrost degradation. Biogeosciences.
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Read PublicationGreenhouse gas emissions from thawing permafrost in arctic ecosystems may amplify global warming, yet estimates of the rate of carbon release, and the proportion of carbon released as methane (CH4) or carbon dioxide (CO2), have a high degree of uncertainty. There are many areas where no measurements exist, and few year-round or long-term records. Existing year-round eddy covariance measurements of arctic CH4 fluxes suggest that nongrowing season emissions make up a significant proportion of tundra systems emissions on an annual basis. Here we present continuous CH4 flux measurements made at Eight Mile Lake, an upland tundra ecosystem undergoing permafrost degradation in Interior Alaska. We found net CH4 emissions throughout the year (1.2 ? 0.011 g C-CH4 m2/yr) that made up 61% of total radiative forcing from annual C emissions (CO2 and CH4; 32.3 g C m2/yr) when taking into account the greenhouse warming potential of CH4 relative to CO2. Nongrowing season emissions accounted for 50% of the annual CH4 budget, characterized by large pulse emissions. These were related to abrupt increases in air and shallow soil temperatures rather than consistent emissions during the zero curtain?a period of the fall/early winter season when subsurface soil temperatures remain near the 0 °C freezing point. Weekly growing season CH4 emissions in 2016 and 2017 were significantly related with thaw depth, and the magnitude of CH4 emissions between these seasons was proportional to the rate of active layer thaw throughout the season.
Terrer C, Vicca S, Stocker BD, Hungate BA, Phillips RP, Reich PB, Finzi AC, Prentice IC (2018) Ecosystem responses to elevated CO2 governed by plant–soil interactions and the cost of nitrogen acquisition. New Phytologist 217(2): 507-522.
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Read PublicationLand ecosystems sequester on average about a quarter of anthropogenic CO2 emissions. It has been proposed that nitrogen (N) availability will exert an increasingly limiting effect on plants’ ability to store additional carbon (C) under rising CO2, but these mechanisms are not well understood. Here, we review findings from elevated CO2 experiments using a plant economics framework, highlighting how ecosystem responses to elevated CO@ may depend on the costs and benefits of plant interactions with mycorrhizal fungi and symbiotic N‐fixing microbes. We found that N‐acquisition efficiency is positively correlated with leaf‐level photosynthetic capacity and plant growth, and negatively with soil C storage. Plants that associate with ectomycorrhizal fungi and N‐fixers may acquire N at a lower cost than plants associated with arbuscular mycorrhizal fungi. However, the additional growth in ectomycorrhizal plants is partly offset by decreases in soil C pools via priming. Collectively, our results indicate that predictive models aimed at quantifying C cycle feedbacks to global change may be improved by treating N as a resource that can be acquired by plants in exchange for energy, with different costs depending on plant interactions with microbial symbionts.
Teubner IE, Forkel M, Jung M, Liu YY, Miralles DG, Parinussa R, van der Schalie R, Vreugdenhil M, Schwalm CR, Tramontana G, Camps-Valls G, Dorigo WA (2018) Assessing the relationship between microwave vegetation optical depth and gross primary production. International Journal of Applied Earth Observation and Geoinformation 65: 79-91.
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Read PublicationAt the global scale, the uptake of atmospheric carbon dioxide by terrestrial ecosystems through photosynthesis is commonly estimated through vegetation indices or biophysical properties derived from optical remote sensing data. Microwave observations of vegetated areas are sensitive to different components of the vegetation layer than observations in the optical domain and may therefore provide complementary information on the vegetation state, which may be used in the estimation of Gross Primary Production (GPP). However, the relation between GPP and Vegetation Optical Depth (VOD), a biophysical quantity derived from microwave observations, is not yet known. This study aims to explore the relationship between VOD and GPP. VOD data were taken from different frequencies (L-, C-, and X-band) and from both active and passive microwave sensors, including the Advanced Scatterometer (ASCAT), the Soil Moisture Ocean Salinity (SMOS) mission, the Advanced Microwave Scanning Radiometer for Earth Observation System (AMSR-E) and a merged VOD data set from various passive microwave sensors. VOD data were compared against FLUXCOM GPP and Solar-Induced chlorophyll Fluorescence (SIF) from the Global Ozone Monitoring Experiment-2 (GOME-2). FLUXCOM GPP estimates are based on the upscaling of flux tower GPP observations using optical satellite data, while SIF observations present a measure of photosynthetic activity and are often used as a proxy for GPP. For relating VOD to GPP, three variables were analyzed: original VOD time series, temporal changes in VOD (ΔVOD), and positive changes in VOD (ΔVOD≥0). Results show widespread positive correlations between VOD and GPP with some negative correlations mainly occurring in dry and wet regions for active and passive VOD, respectively. Correlations between VOD and GPP were similar or higher than between VOD and SIF. When comparing the three variables for relating VOD to GPP, correlations with GPP were higher for the original VOD time series than for ΔVOD or ΔVOD≥0 in case of sparsely to moderately vegetated areas and evergreen forests, while the opposite was true for deciduous forests. Results suggest that original VOD time series should be used jointly with changes in VOD for the estimation of GPP across biomes, which may further benefit from combining active and passive VOD data.
Toda M, Richardson AD (2018) Estimation of plant area index and phenological transition dates from digital repeat photography and radiometric approaches in a hardwood forest in the Northeastern United States. Agricultural and Forest Meteorology 249: 457-466.
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Read PublicationLong-term, continuous digital camera imagery and tower-based radiometric monitoring were conducted at a representative hardwood forest site in the Northeastern United States, part of the AmeriFlux network. In this study, the phenological metrics of the leaf area index (LAI), plant area index (PAI) and associated transition dates (e.g., timing of the onset of leaf expansion and the cessation of leaf fall) were compared using 4-year of data from Bartlett Experimental Forest. We used digital repeat photography (DRP) imagery collected using two different methods (“canopy cover” and “phenocam” approaches), together with above- and below-canopy measurements of photosynthetically active radiation (PAR). The growth-period LAI estimated from canopy cover images (LAICANOPY) and the above and below canopy PAR measurements (LAIfPARt) were within approximately the same range, in term of magnitude, as previous results for multiple comparative methods, although growing-season LAICANOPY was slightly lower (3.11 m2 m−2 to 3.35 m2 m−2) than LAIfPARt (3.19 m2 m−2 to 3.67 m2 m−2). In addition, we derived phenological transition dates from PAICANOPY, PAIfPARt, and color-based metrics calculated from the phenocam imagery (green (GCC) and red (RCC) chromatic coordinates). The transition dates in both spring and autumn differed somewhat according to method, presumably due to the vegetation status detection abilities of each vegetation metric. We found that LAI estimation from canopy cover images may be influenced by automatic exposure settings, which limits the ability to detect subtle changes in phenology during the transition phases in both spring and autumn. Particularly in autumn, the color-based metrics calculated from the phenocam imagery are decoupled from leaf area dynamics and thus PAI. While above and below canopy PAR measurements could yield the better indicators for estimating LAI, its seasonal dynamics, and associated phenological transition dates in long-term monitoring, we argue that there are obvious benefits to the multi-sensor approach used here.
Tong X, Brandt M, Yue Y, Horion S, Wang K, Keersmaecker WD, Tian F, Schurgers G, Xiao X, Luo Y, Chen C, Mynemi R, Shi Z, Chen H, Rensholt R (2018) Increased vegetation growth and carbon stock in China karst via ecological engineering. Nature Sustainability 1(1): 44-50.
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Read PublicationAfforestation and reforestation projects in the karst regions of southwest China aim to combat desertification and improve the ecological environment. However, it remains unclear at what scale conservation efforts have impacted on carbon stocks and if vegetation regrowth occurs at a large spatial scale as intended. Here we use satellite time series data and show a widespread increase in leaf area index (a proxy for green vegetation cover), and aboveground biomass carbon, which contrasted negative trends found in the absence of anthropogenic influence as simulated by an ecosystem model. In spite of drought conditions, aboveground biomass carbon increased by 9% (+0.05 Pg C y−1), mainly in areas of high conservation effort. We conclude that large scale conservation projects can contribute to a greening Earth with positive effects on carbon sequestration to mitigate climate change. At the regional scale, such ecological engineering projects may reduce risks of desertification by increasing the vegetation cover and reducing the ecosystem sensitivity to climate perturbations.
Truettner C, Anderegg WRL, Biondi F, Koch GW, Olge K, Schwalm C, Litvak ME, Shaw JD, Ziaco E (2018) Conifer radial growth response to recent seasonal warming and drought from the southwestern USA. Forest Ecology and Management.
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Read PublicationFuture droughts are expected to become more severe and frequent under future climate change scenarios, likely causing widespread tree mortality in the western USA. Coping with an uncertain future requires an understanding of long-term ecosystem responses in areas where prolonged drought is projected to increase. Tree-ring records are ideally suited for this task. We developed 24 tree-ring chronologies from 20 U.S. Forest Service Forest Inventory and Analysis (FIA) plots in the southwestern USA. Climate variables were derived from the PRISM climate dataset (800-m grid cells) to capture the bimodal precipitation regime of winter snow and summer monsoonal rainfall, as well as warm-season vapor-pressure deficit (VPD) and winter minimum temperature. Based on mixed linear models, radial growth from 1948 to 2013 for four conifer species (Pinus edulis, Juniperus osteosperma, Pinus ponderosa, and Picea engelmannii) responded negatively to warm-season VPD and positively to cold-season precipitation. Pinus spp. benefited from warm-season precipitation linked to the North American monsoon, and Pinus spp. and J. osteosperma radial growth increased with warmer cold-season minimum temperature. However, warmer cold-season minimum temperatures countered the beneficial influence of cold-season precipitation for radial growth in Pinus spp. and J. osteosperma, while P. engelmannii was unaffected. Also, enhanced drying effects of warm-season VPD associated with decreased cold-season precipitation negatively affected radial growth of Pinus spp. and P. engelmannii. Of the four conifer species studied, Pinus spp. are most affected by droughts since 1948, while P. engelmanniiand J. osteosperma appear to be more resilient. Investigating seasonal climate responses and interaction effects on radial growth in areas impacted by severe drought helps identify species that may be particularly at risk from climate change impacts in the Anthropocene.
Trugman AT, Detto M, Bartlett MK, Medvigy D, Anderegg WRL, Schwalm C, Schaffer B, Pacala SW (2018) Tree carbon allocation explains forest drought-kill and recovery patterns. Ecology Letters 21(10): 1552-1560.
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Read PublicationThe mechanisms governing tree drought mortality and recovery remain a subject of inquiry and active debate given their role in the terrestrial carbon cycle and their concomitant impact on climate change. Counter-intuitively, many trees do not die during the drought itself. Indeed, observations globally have documented that trees often grow for several years after drought before mortality. A combination of meta-analysis and tree physiological models demonstrate that optimal carbon allocation after drought explains observed patterns of delayed tree mortality and provides a predictive recovery framework. Specifically, post-drought, trees attempt to repair water transport tissue and achieve positive carbon balance through regrowing drought-damaged xylem. Furthermore, the number of years of xylem regrowth required to recover function increases with tree size, explaining why drought mortality increases with size. These results indicate that tree resilience to drought-kill may increase in the future, provided that CO2 fertilisation facilitates more rapid xylem regrowth.
van Gestel N, Shi Z, van Groenigen KJ, Osenberg CW, Andresen LC, Dukes JS, Hovenden MJ, Luo Y, Michelsen A, Pendall E, Reich PB, Schuur EAG, Hungate BA (2018) Predicting soil carbon loss with warming. Nature 554(7693): E4-E5.
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Read PublicationCrowther et al. Reported that the best predictor of surface soil carbon (top 10 cm) losses in response to warming is the size of the surface carbon stock in the soil (that is, carbon stocks in plots that have not been warmed), finding that soils that are high in soil carbon also lose more carbon under warming conditions. This relationship was based on a linear regression of soil carbon losses and soil carbon stocks in field warming studies, which was then used to project carbon losses over time and to generate a map of soil carbon vulnerability. However, a few extreme data points (high-leverage points) can strongly influence the slope of a regression line. Only 5 of the 49 sites analysed by Crowther et al. are in the upper half of the carbon stock range, which raises the possibility that the relationship they observed could be substantially altered by introducing data from sites with relatively high surface soil carbon stocks.
Walker XJ, Baltzer JL, Cumming SG, Day NJ, Johnstone JF, Rogers BM, Solvik K, Turetsky MR, Mack MC (2018) Soil organic layer combustion in boreal black spruce and jack pine stands of the Northwest Territories, Canada. International Journal of Wildland Fire 27(2): 125-134.
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Read PublicationIncreased fire frequency, extent and severity are expected to strongly affect the structure and function of boreal forest ecosystems. In this study, we examined 213 plots in boreal forests dominated by black spruce (Picea mariana) or jack pine (Pinus banksiana) of the Northwest Territories, Canada, after an unprecedentedly large area burned in 2014. Large fire size is associated with high fire intensity and severity, which would manifest as areas with deep burning of the soil organic layer (SOL). Our primary objectives were to estimate burn depth in these fires and then to characterise landscapes vulnerable to deep burning throughout this region. Here we quantify burn depth in black spruce stands using the position of adventitious roots within the soil column, and in jack pine stands using measurements of burned and unburned SOL depths. Using these estimates, we then evaluate how burn depth and the proportion of SOL combusted varies among forest type, ecozone, plot-level moisture and stand density. Our results suggest that most of the SOL was combusted in jack pine stands regardless of plot moisture class, but that black spruce forests experience complete combustion of the SOL only in dry and moderately well-drained landscape positions. The models and calibrations we present in this study should allow future research to more accurately estimate burn depth in Canadian boreal forests.
Walker XJ, Rogers BM, Baltzer JL, Cumming SG, Day NJ, Goetz SJ, Johnstone JF, Schuur EAG, Turetsky MR, Mack MC (2018) Cross‐scale controls on carbon emissions from boreal forest megafires. Global Change Biology.
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Read PublicationClimate warming and drying is associated with increased wildfire disturbance and the emergence of megafires in North American boreal forests. Changes to the fire regime are expected to strongly increase combustion emissions of carbon (C) which could alter regional C balance and positively feedback to climate warming. In order to accurately estimate C emissions and thereby better predict future climate feedbacks, there is a need to understand the major sources of heterogeneity that impact C emissions at different scales. Here, we examined 211 field plots in boreal forests dominated by black spruce (Picea mariana) or jack pine (Pinus banksiana) of the Northwest Territories (NWT), Canada after an unprecedentedly large area burned in 2014. We assessed both aboveground and soil organic layer (SOL) combustion, with the goal of determining the major drivers in total C emissions, as well as to develop a high spatial resolution model to scale emissions in a relatively understudied region of the boreal forest. On average, 3.35 kg C m−2 was combusted and almost 90% of this was from SOL combustion. Our results indicate that black spruce stands located at landscape positions with intermediate drainage contribute the most to C emissions. Indices associated with fire weather and date of burn did not impact emissions, which we attribute to the extreme fire weather over a short period of time. Using these results, we estimated a total of 94.3 Tg C emitted from 2.85 Mha of burned area across the entire 2014 NWT fire complex, which offsets almost 50% of mean annual net ecosystem production in terrestrial ecosystems of Canada. Our study also highlights the need for fine‐scale estimates of burned area that represent small water bodies and regionally specific calibrations of combustion that account for spatial heterogeneity in order to accurately model emissions at the continental scale.
Wang Y, Ciais P, Goll D, Huang Y, Luo Y, Wang YP, Bloom AA, Broquet G, Hartmann J, Peng S, Penuelas J, Piao S, Sardans J, Stocker BD, Wang R, Zaehle S, Zechmeister-Boltenstern S (2018) GOLUM-CNP v1.0: a data-driven modeling of carbon, nitrogen and phosphorus cycles in major terrestrial biomes. Geoscientific Model Development 11(9):3903-3928.
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Read PublicationGlobal terrestrial nitrogen (N) and phosphorus (P) cycles are coupled to the global carbon (C) cycle for net primary production (NPP), plant C allocation, and decomposition of soil organic matter, but N and P have distinct pathways of inputs and losses. Current C-nutrient models exhibit large uncertainties in their estimates of pool sizes, fluxes, and turnover rates of nutrients, due to a lack of consistent global data for evaluating the models. In this study, we present a new model–data fusion framework called the Global Observation-based Land-ecosystems Utilization Model of Carbon, Nitrogen and Phosphorus (GOLUM-CNP) that combines the CARbon DAta MOdel fraMework (CARDAMOM) data-constrained C-cycle analysis with spatially explicit data-driven estimates of N and P inputs and losses and with observed stoichiometric ratios. We calculated the steady-state N- and P-pool sizes and fluxes globally for large biomes. Our study showed that new N inputs from biological fixation and deposition supplied <span class="inline-formula">>20</span> % of total plant uptake in most forest ecosystems but accounted for smaller fractions in boreal forests and grasslands. New P inputs from atmospheric deposition and rock weathering supplied a much smaller fraction of total plant uptake than new N inputs, indicating the importance of internal P recycling within ecosystems to support plant growth. Nutrient-use efficiency, defined as the ratio of gross primary production (GPP) to plant nutrient uptake, were diagnosed from our model results and compared between biomes. Tropical forests had the lowest N-use efficiency and the highest P-use efficiency of the forest biomes. An analysis of sensitivity and uncertainty indicated that the NPP-allocation fractions to leaves, roots,<span id="page3904"/> and wood contributed the most to the uncertainties in the estimates of nutrient-use efficiencies. Correcting for biases in NPP-allocation fractions produced more plausible gradients of N- and P-use efficiencies from tropical to boreal ecosystems and highlighted the critical role of accurate measurements of C allocation for understanding the N and P cycles.
Wymore AS, Salpas E, Casaburi G, Liu CM, Price LB, Hungate BA, McDowell WH, Marks JC (2018) Effects of plant species on stream bacterial communities via leachate from leaf litter. Hydrobiologia 807(1):131-144.
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Read PublicationLeaf litter provides an important resource to forested stream ecosystems. During leaf fall a significant amount of dissolved organic carbon (DOC) enters streams as leaf leachate. We compared the effects of plant species and leaf leachate bioavailability on the composition of stream bacterial communities and rates of DOC decomposition. We used four common riparian tree species that varied in foliar chemistry, leachate optical properties, and litter decomposition rate. We used laboratory microcosms from two streams and amended with a standard concentration of DOC derived from leaf leachate of the four tree species. After 24 h, we measured rates of DOC biodegradation and determined the composition of the bacterial communities via bar-coded pyrosequencing of the 16S rRNA gene. The composition, diversity, and abundance of the bacterial community differed significantly among plant species from both streams. The phylogenetic distance of the different bacterial communities correlated with species-specific leachate optical properties and rates of DOC biodegradation. Highest rates of DOC decomposition were associated with high tannin and lignin leaf types. Results demonstrate that riparian plant species strongly influences stream bacterial communities via their leachate suggesting that alterations to the presence or abundance of riparian plant taxa may influence these communities and associated ecosystem processes.
Xin Q, Dai Y, Li X, Liu X, Gong P, Richardson AD (2018) A steady-state approximation approach to simulate seasonal leaf dynamics of deciduous broadleaf forests via climate variables. Agricultural and Forest Meteorology 249: 44-56.
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Read PublicationAs leaves are the basic elements of plants that conduct photosynthesis and transpiration, vegetation leaf dynamics controls canopy physical and biogeochemical processes and hence largely influences the interactive exchanges of energy and materials between the land surface and the atmosphere. Given that the processes of plant leaf allocation is highly sensitive to climatological and environmental conditions, developing robust models that simulate leaf dynamics via climate variables contributes a key component to land surface models and coupled land-atmosphere models. Here we propose a new method to simulate seasonal leaf dynamics based on the idea of applying vegetation productivity as a synthesized metric to track and assess the climate suitability to plant growth over time. The method first solves two closed simultaneous equations of leaf phenology and canopy photosynthesis as modeled using the Growing Production-Day model iteratively for deriving the time series of steady-state leaf area index (LAI), and then applies the method of simple moving average to account for the time lagging of leaf allocation behind steady-state LAI. The seasonal LAI simulated using the developed method agree with field measurements from a selection of AmeriFlux sites as indicated by high coefficient of determination (R2 = 0.801) and low root mean square error (RMSE = 0.924 m2/m2) and with satellite-derived data (R2 = 0.929 and RMSE = 0.650 m2/m2) for the studied flux tower sites. Moreover, the proposed method is able to simulate seasonal LAI of deciduous broadleaf forests that match with satellite-derived LAI time series across the entire eastern United States. Comparative modeling studies suggest that the proposed method produces more accurate results than the method based on Growing Season Index in terms of correlation coefficients and error metrics. The developed method provides a complete solution to modeling seasonal leaf dynamics as well as canopy productivity solely using climate variables.
Xu H, Zhang T, Luo Y, Huang X, Xue W (2018) Parameter calibration in global soil carbon models using surrogate-based optimization. Geoscientific Model Development 11(7): 3027-3044.
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Read PublicationSoil organic carbon (SOC) has a significant effect on carbon emissions and climate change. However, the current SOC prediction accuracy of most models is very low. Most evaluation studies indicate that the prediction error mainly comes from parameter uncertainties, which can be improved by parameter calibration. Data assimilation techniques have been successfully employed for the parameter calibration of SOC models. However, data assimilation algorithms, such as the sampling-based Bayesian Markov chain Monte Carlo (MCMC), generally have high computation costs and are not appropriate for complex global land models. This study proposes a new parameter calibration method based on surrogate optimization techniques to improve the prediction accuracy of SOC. Experiments on three types of soil carbon cycle models, including the Community Land Model with the Carnegie–Ames–Stanford Approach biogeochemistry submodel (CLM-CASA') and two microbial models show that the surrogate-based optimization method is effective and efficient in terms of both accuracy and cost. Compared to predictions using the tuned parameter values through Bayesian MCMC, the root mean squared errors (RMSEs) between the predictions using the calibrated parameter values with surrogate-base optimization and the observations could be reduced by up to 12% for different SOC models. Meanwhile, the corresponding computational cost is lower than other global optimization algorithms.
Yuan MM, Zhang J, Xue K, Wu L, Deng Y, Deng J, Hale L, Zhou X, He Z, Yang Y, Van Nostrand JD, Schuur EAG, Konstantinidis KT, Penton CR, Cole JR, Tiedje JM, Luo Y, Zhou J (2018) Microbial functional diversity covaries with permafrost thaw‐induced environmental heterogeneity in tundra soil. Global Change Biology 24(1): 297-307.
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Read PublicationPermafrost soil in high latitude tundra is one of the largest terrestrial carbon (C) stocks and is highly sensitive to climate warming. Understanding microbial responses to warming‐induced environmental changes is critical to evaluating their influences on soil biogeochemical cycles. In this study, a functional gene array (i.e., geochip 4.2) was used to analyze the functional capacities of soil microbial communities collected from a naturally degrading permafrost region in Central Alaska. Varied thaw history was reported to be the main driver of soil and plant differences across a gradient of minimally, moderately, and extensively thawed sites. Compared with the minimally thawed site, the number of detected functional gene probes across the 15–65 cm depth profile at the moderately and extensively thawed sites decreased by 25% and 5%, while the community functional gene β‐diversity increased by 34% and 45%, respectively, revealing decreased functional gene richness but increased community heterogeneity along the thaw progression. Particularly, the moderately thawed site contained microbial communities with the highest abundances of many genes involved in prokaryotic C degradation, ammonification, and nitrification processes, but lower abundances of fungal C decomposition and anaerobic‐related genes. Significant correlations were observed between functional gene abundance and vascular plant primary productivity, suggesting that plant growth and species composition could be co‐evolving traits together with microbial community composition. Altogether, this study reveals the complex responses of microbial functional potentials to thaw‐related soil and plant changes and provides information on potential microbially mediated biogeochemical cycles in tundra ecosystems.
Zhang T, Luo Y, Chen HYH, Ruan H (2018) Responses of litter decomposition and nutrient release to N addition: A meta-analysis of terrestrial ecosystems. Applied Soil Ecology 128: 35-42.
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Read PublicationAs atmospheric nitrogen (N) concentrations increase, it can wreak havoc on the entire planet, as well as the fragile ecosystems, once it exceeds the demand of ecosystems. Chronically elevated N deposition affects litter decomposition, which is a crucial process that controls nutrient cycling, soil fertility, and primary productivity. Nevertheless, the responses of litter decomposition and nutrient release to N addition remain elusive. Here we conduct a meta-analysis using 3434 paired observations from 55 publications to evaluate these responses. We found that although litter decomposition rate did not change significantly under N addition when averaged across all studies, it decreased with N application rate and experimental duration, showing that it was stimulated at low levels but suppressed at high levels of N application and duration. Phosphorus released more slowly under N enrichment, and this response became greater with longer duration. Moreover, the decomposition of lignin was depressed under N addition, and this effect was more pronounced with the increase of N application rate and experimental duration. Importantly, in terms of different ecosystems, the decomposition of litter was significantly inhibited by N addition in plantations, but was promoted in secondary forests, and there were no significant changes in primary forests, grasslands and wetlands. The responses of litter mass loss, along with the release of nutrients to N fertilization, changed with mean annual temperature and mean annual precipitation of the study sites. Our results provided a synthetic understanding of the effects of N addition on the decomposition of litter and nutrient release under climate change scenarios.
Zhang X, Jayavelu S, Liu L, Friedl MA, Henebry GM, Liu Y, Schaaf CB, Richardson AD, Gray J (2018) Evaluation of land surface phenology from VIIRS data using time series of PhenoCam imagery. Agricultural and Forest Meteorology 256-257: 137-149.
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Read PublicationLand surface phenology (LSP) has been widely retrieved from time series of various satellite instruments in order to monitor climate change and ecosystem dynamics. However, any evaluation of the quality of LSP data sets is quite challenging because the in situ observations on a limited number of individual trees, shrubs, or other plants are rarely representative of the landscape sampled in a single satellite pixel. Moreover, vegetation indices detecting biophysical features of vegetation seasonality are different from (but related to) the specific plant life history stages observed by humans at ground level. This study is the first comprehensive evaluation of the LSP product derived from Visible Infrared Imaging Radiometer Suite (VIIRS) data using both MODIS LSP products and observations from the PhenoCam network across the Contiguous United States during 2013 and 2014. PhenoCam observes vegetation canopy over a landscape at very high frequency, providing nearly continuous canopy status and enabling the estimate of discrete phenophase using vegetation indices that are conceptually similar to satellite data. Six phenological dates (greenup onset, mid-greenup phase, maturity onset, senescence onset, mid-senescence phase, and dormancy onset) were retrieved separately from daily VIIRS NDVI (normalized difference vegetative index) and EVI2 (two-band enhanced vegetation index) time series. Similarly, the six phenological dates were also extracted from the 30-min time series of PhenoCam data using GCC (green chromatic coordinate) and VCI (vegetation contrast index) separately. Phenological dates derived from VIIRS NDVI and EVI2 and PhenoCam GCC and VCI were generally comparable for the vegetation greenup phase, but differed considerably for the senescence phase. Although all indices captured green leaf development effectively, performance discrepancies arose due to their ability to track the mixture of senescing leaf colors. PhenoCam GCC and VCI phenological observations were in better agreement with the phenological dates from VIIRS EVI2 than from VIIRS NDVI. Further, the VIIRS EVI2 phenological metrics were more similar to those from PhenoCam VCI than from PhenoCam GCC time series. Overall, the average absolute difference between the VIIRS EVI2 and PhenoCam VCI phenological dates was 7–11 days in the greenup phase and 10–13 days in the senescence phase. The difference was smaller in forests, followed by grasslands and croplands, and then savannas. Finally, the phenological dates derived from VIIRS EVI2 were compared with MODIS detections, which showed a good agreement with an average absolute difference less than a week except for the senescence onset. These results for the first time demonstrate the upper boundary of uncertainty in VIIRS LSP detections and the continuity to MODIS LSP product.
Zhou S, Liang J, Lu X, Li Q, Jiang L, Zhang Y, Schwalm CR, Fisher JB, Tjiputra J, Sitch S, Ahlström A, Huntzinger DN, Huang Y, Wang G, Luo Y (2018) Sources of uncertainty in modeled land carbon storage within and across three MIPs: Diagnosis with three new techniques. Journal of Climate 31(7): 2833-2851.
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Read PublicationTerrestrial carbon cycle models have incorporated increasingly more processes as a means to achieve more-realistic representations of ecosystem carbon cycling. Despite this, there are large across-model variations in the simulation and projection of carbon cycling. Several model intercomparison projects (MIPs), for example, the fifth phase of the Coupled Model Intercomparison Project (CMIP5) (historical simulations), Trends in Net Land?Atmosphere Carbon Exchange (TRENDY), and Multiscale Synthesis and Terrestrial Model Intercomparison Project (MsTMIP), have sought to understand intermodel differences. In this study, the authors developed a suite of new techniques to conduct post-MIP analysis to gain insights into uncertainty sources across 25 models in the three MIPs. First, terrestrial carbon storage dynamics were characterized by a three-dimensional (3D) model output space with coordinates of carbon residence time, net primary productivity (NPP), and carbon storage potential. The latter represents the potential of an ecosystem to lose or gain carbon. This space can be used to measure how and why model output differs. Models with a nitrogen cycle generally exhibit lower annual NPP in comparison with other models, and mostly negative carbon storage potential. Second, a transient traceability framework was used to decompose any given carbon cycle model into traceable components and identify the sources of model differences. The carbon residence time (or NPP) was traced to baseline carbon residence time (or baseline NPP related to the maximum carbon input), environmental scalars, and climate forcing. Third, by applying a variance decomposition method, the authors show that the intermodel differences in carbon storage can be mainly attributed to the baseline carbon residence time and baseline NPP (>90% in the three MIPs). The three techniques developed in this study offer a novel approach to gain more insight from existing MIPs and can point out directions for future MIPs. Since this study is conducted at the global scale for an overview on intermodel differences, future studies should focus more on regional analysis to identify the sources of uncertainties and improve models at the specified mechanism level.
Zhou X, Xu X, Zhou G, Luo Y (2018) Temperature sensitivity of soil organic carbon decomposition increased with mean carbon residence time: Field incubation and data assimilation. Global Change Biology 24(2): 810-822.
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Read PublicationTemperature sensitivity of soil organic carbon (SOC) decomposition is one of the major uncertainties in predicting climate‐carbon (C) cycle feedback. Results from previous studies are highly contradictory with old soil C decomposition being more, similarly, or less sensitive to temperature than decomposition of young fractions. The contradictory results are partly from difficulties in distinguishing old from young SOC and their changes over time in the experiments with or without isotopic techniques. In this study, we have conducted a long‐term field incubation experiment with deep soil collars (0–70 cm in depth, 10 cm in diameter of PVC tubes) for excluding root C input to examine apparent temperature sensitivity of SOC decomposition under ambient and warming treatments from 2002 to 2008. The data from the experiment were infused into a multi‐pool soil C model to estimate intrinsic temperature sensitivity of SOC decomposition and C residence times of three SOC fractions (i.e., active, slow, and passive) using a data assimilation (DA) technique. As active SOC with the short C residence time was progressively depleted in the deep soil collars under both ambient and warming treatments, the residences times of the whole SOC became longer over time. Concomitantly, the estimated apparent and intrinsic temperature sensitivity of SOC decomposition also became gradually higher over time as more than 50% of active SOC was depleted. Thus, the temperature sensitivity of soil C decomposition in deep soil collars was positively correlated with the mean C residence times. However, the regression slope of the temperature sensitivity against the residence time was lower under the warming treatment than under ambient temperature, indicating that other processes also regulated temperature sensitivity of SOC decomposition. These results indicate that old SOC decomposition is more sensitive to temperature than young components, making the old C more vulnerable to future warmer climate.
Zhou Z, Wang C, Luo Y (2018) Response of soil microbial communities to altered precipitation: A global synthesis. Global Ecology and Biogeography 27(9): 1121-1136.
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Read PublicationClimate change intensifies the hydrological cycle and consequently alters precipitation regimes. Accurately assessing future carbon (C) budgets depends on understanding the influence of altered precipitation on both aboveground C cycling and belowground processes. Our goal was to explore generalities and mechanisms of responses of soil microbial communities to altered precipitation and implications for C cycling in terrestrial ecosystems. Location Global. Time period 2001?2017. Major taxa studied Soil microbes. Methods We used the meta-analytical technique to synthesize data of 41 increased (IPPT) and 53 decreased precipitation (DPPT) studies from 65 publications worldwide. The data covered broad variations in climate, percentage of precipitation change, experimental duration and soil properties. Results The fungi to bacteria ratio did not show a water-tolerant shift, but the community compositions within the bacteria did. Microbial biomass showed a higher response to moderate IPPT than moderate DPPT, whereas it was more sensitive to extreme DPPT than extreme IPPT, suggesting that the responses of microbial biomass to altered precipitation are double asymmetric. However, such asymmetric responses of microbial biomass varied with climate humidity and soil texture: microbial biomass was more sensitive to IPPT at xeric sites than at mesic sites, whereas it was more responsive to DPPT in humid areas; microbial biomass in coarse-textured soils was more sensitive to altered precipitation than that in fine-textured soils. In addition, microbial response was positively correlated with the responses of aboveground/belowground plant biomass, soil respiration and organic C content. Main conclusions Our meta-analysis provides the first evidence that the asymmetric response of microbial biomass to altered precipitation varies with climate humidity and soil texture. Given the coordinated responses in the plant?soil?microorganism C continuum, our synthesis extends the double asymmetric model and provides a framework for understanding and modelling responses of ecosystem C cycling to global precipitation change.
Zou J, Tobin B, Luo Y, Osborne B (2018) Differential responses of soil CO2 and N2O fluxes to experimental warming. Agricultural and Forest Meteorology 259: 11-22.
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Read PublicationLand-use conversions and elevated temperature can impact on carbon dioxide (CO2) and nitrous oxide (N2O) emissions, both of which are important greenhouse gasses (GHGs). Afforestation activity has increased significantly over the last century with a significant focus in recent years directed at offsetting GHG emissions, as forests have a large capacity to store carbon (C) and nitrogen (N) as well as affecting CO2 and N2O emissions. However, the impact of warming on GHG offsetting is unclear. This study was conducted in a forest and a grassland to investigate the effect of afforestation and warming, using infrared heaters, on soil fluxes of CO2 and N2O. Warming significantly increased the daily mean soil temperatures at a depth of 5 cm by 1.7 °C and reduced the soil moisture by ∼5% in the forest from March 2014 to February 2016. In the grassland, there were no significant increases in temperature and moisture with warming and no impact on the soil fluxes of CO2 and N2O. In the forest, elevated soil temperature enhanced the average soil CO2 efflux by 23% but had no effect on soil N2O fluxes. Warming decreased the temperature sensitivity by 13% and 23% at the forest and grassland, respectively. The soil fluxes of CO2 increased exponentially with temperature and decreased linearly with the reduction in soil moisture, and were much larger in the grassland compared to the forest. However the grassland proved to be a larger sink for N2O than the forest. Irrespective of warming treatments, all measured pools were significantly larger in the grassland compared to the forest. Our results imply that afforestation may have a bigger effect than warming on soil CO2 and N2O fluxes within the range of temperatures used and that afforestation dramatically lowers the inorganic, organic and microbial C and N pools, that could, in turn, impact on the responses of forest soils to future global warming.
Zou J, Tobin B, Luo Y, Osborne B (2018) Response of soil respiration and its components to experimental warming and water addition in a temperate Sitka spruce forest ecosystem. Agricultural and Forest Meteorology 260-261: 204-215.
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Read PublicationFuture climate change is expected to alter the terrestrial carbon cycle through its impact on soil respiration. In this study, we determined the responses of soil respiration and its components to experimental warming with or without water addition. A replicated <em>in situ</em> heating (∼2 °C above ambient soil temperatures) and water addition (170 mm in total each year) experiment was carried out for the first time in a temperate plantation forest of Sitka spruce over the period 2014–2016. R<sub>h</sub>was measured inside deep collars (35 cm deep) that excluded root growth, while R<sub>s</sub>was measured using the static chamber approach and near-surface collars (5 cm deep) and R<sub>a</sub> calculated by subtracting R<sub>h</sub> from total soil respiration (R<sub>s</sub>). Experimental warming significantly increased R<sub>s</sub>, R<sub>h</sub> and R<sub>h</sub>/R<sub>s</sub>, but had no effect on R<sub>a</sub>. In contrast, none of the respiration components were affected by water addition. Warming increased annual R<sub>h</sub> by 62% but had no effect on R<sub>a</sub>. Overall, warming did not significantly increase annual R<sub>s</sub>. Warming showed a stronger impact on R<sub>s</sub> in the non-growing season but had a smaller impact in the growing season. Warming increased R<sub>a</sub> in the non-growing season but decreased it in the growing season. The effects of warming on R<sub>h</sub> were similar for the two periods. Our results highlight the differential response of R<sub>a</sub> and R<sub>h</sub> to warming, which was mediated by water addition or season. For this and other similar forest sites that don’t experience water limitation, global warming may have a positive feedback on atmospheric CO<sub>2</sub>concentrations through enhanced soil respiration.
2017
Alexander HD, Mack MC (2017) Gap regeneration within mature deciduous forests of Interior Alaska: Implications for future forest change. Forest Ecology and Management 396: 35-43.
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Read PublicationIncreased fire severity in boreal forests of Interior Alaska is shifting forest canopy composition from black spruce (Picea mariana) to deciduous species, including trembling aspen (Populus tremuloides) and Alaska paper birch (Betula neoalaskana). Because deciduous trees are less flammable than black spruce, the dominant disturbance regime in deciduous forests could move away from fire to one of gap disturbances. In this study, we quantified forest gap characteristics and vegetation within eight mature (62-119-yr-old) deciduous stands in Interior Alaska. Canopy gaps were generally small (true gap area <50 m(2)), formed by the mortality of 4-16 gap makers (which were always deciduous trees), and occupied similar to 17-29% of the forest except in the oldest stand, where gap fraction exceeded 45%, and in one anomalous 84-yr old stand, where gaps were absent. Canopy openness increased linearly with gap area, but density of both deciduous and evergreen tree recruits was generally low and insufficient to create future stands with densities similar to those currently found in mature stands across the landscape. Canopy openness was instead correlated with decreased leaf litter cover and increased cover of moss, lichen, and evergreen shrubs. Given the low recruitment of trees with canopy gaps and the decreased probability of fire, deciduous stands will likely "transition to non-forested areas or low density stands once overstory trees reach maturity and die. This could have numerous implications for ecosystem function, including carbon (C), water, and energy balance, and potential feedbacks to future fire occurrence and regional climate. (C) 2017 Elsevier B.V. All rights reserved.
Brown LA, Dash J, Ogutu BO, Richardson AD (2017) On the relationship between continuous measures of canopy greenness derived using near-surface remote sensing and satellite-derived vegetation products. Agricultural and Forest Meteorology 247: 280-292.
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Read PublicationOver the last two decades, satellite-derived estimates of biophysical variables have been increasingly used in operational services, requiring quantification of their accuracy and uncertainty. Evaluating satellite-derived vegetation products is challenging due to their moderate spatial resolution, the heterogeneity of the terrestrial landscape, and difficulties in adequately characterising spatial and temporal vegetation dynamics. In recent years, near-surface remote sensing has emerged as a potential source of data against which satellite-derived vegetation products can be evaluated. Several studies have focussed on the evaluation of satellite-derived phenological transition dates, however in most cases the shape and magnitude of the underlying time-series are neglected. In this paper, we investigated the relationship between the green chromatic coordinate (GCC) derived using near-surface remote sensing and a range of vegetation products derived from the Medium Resolution Imaging Spectrometer (MERIS) throughout the growing season. Moderate to strong relationships between the GCC and vegetation products derived from MERIS were observed at deciduous forest sites. Weak relationships were observed over evergreen forest sites as a result of their subtle seasonality, which is likely masked by atmospheric, bidirectional reflectance distribution function (BRDF), and shadowing effects. Temporal inconsistencies were attributed to the oblique viewing geometry of the digital cameras and differences in the incorporated spectral bands. In addition, the commonly observed summer decline in GCC values was found to be primarily associated with seasonal variations in brown pigment concentration, and to a lesser extent illumination geometry. At deciduous sites, increased sensitivity to initial increases in canopy greenness was demonstrated by the GCC, making it particularly well-suited to identifying the start of season when compared to satellite-derived vegetation products. Nevertheless, in some cases, the relationship between the GCC and vegetation products derived from MERIS was found to saturate asymptotically. This limits the potential of the approach for evaluation of the vegetation products that underlie satellite-derived phenological transition dates, and for the continuous monitoring of vegetation during the growing season, particularly at medium to high biomass study sites.
Celis G, Mauritz M, Bracho R, Salmon VG, Webb EE, Hutchings J, Natali SM, Schädel C, Crummer KG, Schuur EAG (2017) Tundra is a consistent source of CO2 at a site with progressive permafrost thaw during six years of chamber and eddy covariance measurements. Biogeosciences.
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Read PublicationCurrent and future warming of high latitude ecosystems will play an important role in climate change through feedbacks to the global carbon cycle. This study compares six years of CO2 flux measurements in moist acidic tundra using autochambers and eddy covariance (Tower) approaches. We found that the tundra was an annual source of CO2 to the atmosphere as indicated by net ecosystem exchange using both methods with a combined mean of 105 +/- 17 g CO2 - C m-2 y-1 across methods and years (Tower 87 +/- 17 and Autochamber 123 +/- 14). The difference between methods was largest early in the observation period, with Autochambers indicated a greater CO2 source to the atmosphere. This discrepancy diminished through time and in the final year the Autochambers measured a greater sink strength than tower. Active layer thickness (ALT) was a significant driver of NEE, GPP, and R eco and could account for differences between Autochamber and Tower. The stronger source initially was attributed lower summer season gross primary production (GPP) during the first three years, coupled with lower ecosystem respiration (R eco) during the first year. The combined suppression of GPP and R eco in the first year of Autochamber measurements could be the result of the experimental setup. Root damage associated with Autochamber soil collar installation may have lowered the plant community's capacity to fix C, but recovered within three years. While this ecosystem was a consistent CO2 sink during the summer, CO2 emissions during the non-summer months offset summer CO2 uptake each year.
D'Antonio CM, Yelenik SG, Mack MC (2017) Ecosystem vs. community recovery 25 years after grass invasions and fire in a subtropical woodland. Journal Of Ecology 105(6): 1462-1474.
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Read PublicationDespite a large body of research documenting invasive plant impacts, few studies have followed individual invaded sites over decades to observe how they change, and none have contrasted how compositional impacts from invasion compare to ecosystem-process impacts over a multi-decadal time-scale.
Using direct measurements of plant density and composition and of ecosystems processes, we evaluate how ecosystem structure, above-ground net primary production (ANPP), and above-ground and soil nutrient pools compare over 25 years since fire and C4 grass invasions disrupted seasonally dry Hawaiian woodlands. We compare structure and function between primary woodland that has never burned and is largely native species-dominated, with sites that had been the same woodland type but burned in alien-grass-fuelled fires in the 1970s and 1980s. The sites have not experienced fires since 1987.
We report here that woody plant composition and structure continue to be dramatically changed by the initial invasions and fires that occurred 25 years ago and invaders continue to dominate in burned sites. This is reflected in continued low plant carbon pools in burned compared to unburned sites. Yet ANPP and N storage, which were dramatically lower in the initial decade after invasive-grass fuelled fires, have increased and are now indistinguishable from values measured in intact woodlands. Soil carbon pools were resilient to both invasion and fire initially and over time.
Above-ground net primary production has recovered because of invasion of burned sites by a non-native N-fixing tree rather than because of recovery of native species. This invasive N-fixing tree is unlikely to return C storage of the invaded sites to those of unburned woodland because of its tissue and growth characteristics and its interactions with invasive grasses. It does not facilitate native species but rather promotes a persistent invasive grass/N-fixer savanna.
Synthesis. We conclude that fire, an unusual disturbance in this system, has perpetuated the dominance of these sites by invasive species and that despite the dramatic recovery of above-ground net primary production and N pools, the ecosystem continues to be in a distinctly different state than the pre-fire, pre-Melinis community. Thus, despite the absence of further disturbance (fire), there is no evidence that succession towards the original ecosystem is occurring. The fact that N pools and above-ground net primary production recover because of a new invader (Morella faya), highlights the unpredictability of ecosystem trajectories in the face of altered regional species pools.
Fang Y, Michalak AM, Schwalm CR, Huntzinger DN, Berry JA, Ciais P, Piao S, Poulter B, Fisher JB, Cook RB, Hayes D, Huang M, Ito A, Jain A, Lei H, Lu C, Mao J, Parazoo NC, Peng S, Ricciuto DM, Shi X, Tao B, Tian H, Wang W, Wei Y and Yang J (2017) Global land carbon sink response to temperature and precipitation varies with ENSO phase. Environmental Research Letters 12(6).
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Read PublicationClimate variability associated with the El Niño-Southern Oscillation (ENSO) and its consequent impacts on land carbon sink interannual variability have been used as a basis for investigating carbon cycle responses to climate variability more broadly, and to inform the sensitivity of the tropical carbon budget to climate change. Past studies have presented opposing views about whether temperature or precipitation is the primary factor driving the response of the land carbon sink to ENSO. Here, we show that the dominant driver varies with ENSO phase. Whereas tropical temperature explains sink dynamics following El Niño conditions (r TG,P = 0.59, p < 0.01), the post La Niña sink is driven largely by tropical precipitation (r PG,T = −0.46, p = 0.04). This finding points to an ENSO-phase-dependent interplay between water availability and temperature in controlling the carbon uptake response to climate variations in tropical ecosystems. We further find that none of a suite of ten contemporary terrestrial biosphere models captures these ENSO-phase-dependent responses, highlighting a key uncertainty in modeling climate impacts on the future of the global land carbon sink.
Feng W, Liang J, Hale LE, Jung CG, Chen J, Zhou Z, Xu M, Yuan M, Wu L, Bracho R, Pegoraro E, Schuur EAG, Luo Y (2017) Enhanced decomposition of stable soil organic carbon and microbial catabolic potentials by long‐term field warming. Global Change Biology 23(11): 4765-4776.
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Read PublicationQuantifying soil organic carbon (SOC) decomposition under warming is critical to predict carbon–climate feedbacks. According to the substrate regulating principle, SOC decomposition would decrease as labile SOC declines under field warming, but observations of SOC decomposition under warming do not always support this prediction. This discrepancy could result from varying changes in SOC components and soil microbial communities under warming. This study aimed to determine the decomposition of SOC components with different turnover times after subjected to long‐term field warming and/or root exclusion to limit C input, and to test whether SOC decomposition is driven by substrate lability under warming. Taking advantage of a 12‐year field warming experiment in a prairie, we assessed the decomposition of SOC components by incubating soils from control and warmed plots, with and without root exclusion for 3 years. We assayed SOC decomposition from these incubations by combining inverse modeling and microbial functional genes during decomposition with a metagenomic technique (GeoChip). The decomposition of SOC components with turnover times of years and decades, which contributed to 95% of total cumulative CO2 respiration, was greater in soils from warmed plots. But the decomposition of labile SOC was similar in warmed plots compared to the control. The diversity of C‐degradation microbial genes generally declined with time during the incubation in all treatments, suggesting shifts of microbial functional groups as substrate composition was changing. Compared to the control, soils from warmed plots showed significant increase in the signal intensities of microbial genes involved in degrading complex organic compounds, implying enhanced potential abilities of microbial catabolism. These are likely responsible for accelerated decomposition of SOC components with slow turnover rates. Overall, the shifted microbial community induced by long‐term warming accelerates the decomposition of SOC components with slow turnover rates and thus amplify the positive feedback to climate change.
Hungate BA, Barbier BE, Ando AW, Marks SP, Reich PB, van Gestel N, Tilman D, Knops JM, Hooper DU, Butterfield BJ, Cardinale BJ (2017) The economic value of grassland species for carbon storage. Science Advances 3(4).
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Read PublicationCarbon storage by ecosystems is valuable for climate protection. Biodiversity conservation may help increase carbon storage, but the value of this influence has been difficult to assess. We use plant, soil, and ecosystem carbon storage data from two grassland biodiversity experiments to show that greater species richness increases economic value: Increasing species richness from 1 to 10 had twice the economic value of increasing species richness from 1 to 2. The marginal value of each additional species declined as species accumulated, reflecting the nonlinear relationship between species richness and plant biomass production. Our demonstration of the economic value of biodiversity for enhancing carbon storage provides a foundation for assessing the value of biodiversity for decisions about land management. Combining carbon storage with other ecosystem services affected by biodiversity may well enhance the economic arguments for conservation even further.
Jean M, Alexander HD, Mack MC, Johnstone JF (2017) Patterns of bryophyte succession in a 160 year chronosequence in deciduous and coniferous forests of boreal Alaska. Canadian Journal of Forest Research 47(8): 1021-1032.
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Read PublicationBryophytes are dominant components of boreal forest understories and play a large role in regulating soil microclimate and nutrient cycling. Therefore, shifts in bryophyte communities have the potential to affect boreal forests’ ecosystem processes. We investigated how bryophyte communities varied in 83 forest stands in interior Alaska that ranged in age (since fire) from 8 to 163 years and had canopies dominated by deciduous broadleaf (Populus tremuloides Michx. or Betula neoalaskana Sarg.) or coniferous trees (Picea mariana Mill B.S.P.). In each stand, we measured bryophyte community composition, along with environmental variables (e.g., organic layer depth, leaf litter cover, moisture). Bryophyte communities were initially similar in deciduous vs. coniferous forests but diverged in older stands in association with changes in organic layer depth and leaf litter cover. Our data suggest two tipping points in bryophyte succession: one at the disappearance of early colonizing taxa 20 years after fire and another at 40 years after fire, which corresponds to canopy closure and differential leaf litter inputs in mature deciduous and coniferous canopies. Our results enhance understanding of the processes that shape compositional patterns and ecosystem services of bryophytes in relation to stand age, canopy composition, and changing disturbances such as fire that may trigger changes in canopy composition.
Jeffery S, Abalos D, Prodana M, Bastos A, van Groenigen JW, Hungate BA, Verheijen F (2017) Biochar boosts tropical but not temperate crop yields. Environmental Research Letters.
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Read PublicationApplying biochar to soil is thought to have multiple benefits, from helping mitigate climate change [1, 2], to managing waste [3] to conserving soil [4]. Biochar is also widely assumed to boost crop yield [5, 6], but there is controversy regarding the extent and cause of any yield benefit [7]. Here we use a global-scale meta-analysis to show that biochar has, on average, no effect on crop yield in temperate latitudes, yet elicits a 25% average increase in yield in the tropics. In the tropics, biochar increased yield through liming and fertilization, consistent with the low soil pH, low fertility, and low fertilizer inputs typical of arable tropical soils. We also found that, in tropical soils, high-nutrient biochar inputs stimulated yield substantially more than low-nutrient biochar, further supporting the role of nutrient fertilization in the observed yield stimulation. In contrast, arable soils in temperate regions are moderate in pH, higher in fertility, and generally receive higher fertilizer inputs, leaving little room for additional benefits from biochar. Our findings demonstrate that the yield-stimulating effects of biochar are not universal, but may especially benefit agriculture in low-nutrient, acidic soils in the tropics. Biochar management in temperate zones should focus on potential non-yield benefits such as lime and fertilizer cost savings, greenhouse gas emissions control, and other ecosystem services.
Jiang L, Shi Z, Xia J, Liang J, Lu X, Wang Y, Luo Y (2017) Transient traceability analysis of land carbon storage dynamics: Procedures and its application to two forest ecosystems. Journal of Advances in Modeling Earth Systems 9(8): 2822-2835.
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Read PublicationUptake of anthropogenically emitted carbon (C) dioxide by terrestrial ecosystem is critical for determining future climate. However, Earth system models project large uncertainties in future C storage. To help identify sources of uncertainties in model predictions, this study develops a transient traceability framework to trace components of C storage dynamics. Transient C storage (X) can be decomposed into two components, C storage capacity (Xc) and C storage potential (Xp). Xc is the maximum C amount that an ecosystem can potentially store and Xp represents the internal capacity of an ecosystem to equilibrate C input and output for a network of pools. Xc is codetermined by net primary production (NPP) and residence time (τN), with the latter being determined by allocation coefficients, transfer coefficients, environmental scalar, and exit rate. Xp is the product of redistribution matrix (τch) and net ecosystem exchange. We applied this framework to two contrasting ecosystems, Duke Forest and Harvard Forest with an ecosystem model. This framework helps identify the mechanisms underlying the responses of carbon cycling in the two forests to climate change. The temporal trajectories of X are similar between the two ecosystems. Using this framework, we found that different mechanisms lead to a similar trajectory between the two ecosystems. This framework has potential to reveal mechanisms behind transient C storage in response to various global change factors. It can also identify sources of uncertainties in predicted transient C storage across models and can therefore be useful for model intercomparison.
Jiang Y, van Groenigen KJ, Haung S, Hungate BA, van Kessel C, Hu S, Zhang J, Wu L, Yan X, Wang L, Chen J, Hang X, Zhang Y, Horwath WR, Ye R, Linquist BA, Song Z, Zheng C, Deng A, Zhang W (2017) Higher yields and lower methane emissions with new rice cultivars. Global Change Biology 23(11): 4728-4738.
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Read PublicationBreeding high‐yielding rice cultivars through increasing biomass is a key strategy to meet rising global food demands. Yet, increasing rice growth can stimulate methane (CH4) emissions, exacerbating global climate change, as rice cultivation is a major source of this powerful greenhouse gas. Here, we show in a series of experiments that high‐yielding rice cultivars actually reduce CH4 emissions from typical paddy soils. Averaged across 33 rice cultivars, a biomass increase of 10% resulted in a 10.3% decrease in CH4 emissions in a soil with a high carbon (C) content. Compared to a low‐yielding cultivar, a high‐yielding cultivar significantly increased root porosity and the abundance of methane‐consuming microorganisms, suggesting that the larger and more porous root systems of high‐yielding cultivars facilitated CH4 oxidation by promoting O2 transport to soils. Our results were further supported by a meta‐analysis, showing that high‐yielding rice cultivars strongly decrease CH4 emissions from paddy soils with high organic C contents. Based on our results, increasing rice biomass by 10% could reduce annual CH4 emissions from Chinese rice agriculture by 7.1%. Our findings suggest that modern rice breeding strategies for high‐yielding cultivars can substantially mitigate paddy CH4 emission in China and other rice growing regions.
Jung M, Reichstein M, Schwalm CR, Huntingford C, Sitch S, Ahlström A, Arneth A, Camps-Valls G, Ciais P, Friedlingstein P, Gans F, Ichii K, Jain AK, Kato E, Papale D, Poulter B, Raduly B, Rödenbeck C, Tramontana G, Viovy N, Wang YP, Weber U, Zaehle S & Zeng N (2017) Compensatory water effects link yearly global land CO2 sink changes to temperature. Nature 541: 516–520.
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Read PublicationLarge interannual variations in the measured growth rate of atmospheric carbon dioxide (CO2) originate primarily from fluctuations in carbon uptake by land ecosystems. It remains uncertain, however, to what extent temperature and water availability control the carbon balance of land ecosystems across spatial and temporal scales. Here we use empirical models based on eddy covariance data and process-based models to investigate the effect of changes in temperature and water availability on gross primary productivity (GPP), terrestrial ecosystem respiration (TER) and net ecosystem exchange (NEE) at local and global scales. We find that water availability is the dominant driver of the local interannual variability in GPP and TER. To a lesser extent this is true also for NEE at the local scale, but when integrated globally, temporal NEE variability is mostly driven by temperature fluctuations. We suggest that this apparent paradox can be explained by two compensatory water effects. Temporal water-driven GPP and TER variations compensate locally, dampening water-driven NEE variability. Spatial water availability anomalies also compensate, leaving a dominant temperature signal in the year-to-year fluctuations of the land carbon sink. These findings help to reconcile seemingly contradictory reports regarding the importance of temperature and water in controlling the interannual variability of the terrestrial carbon balance. Our study indicates that spatial climate covariation drives the global carbon cycle response.
Kerhoulas LP, Kolb TE, Koch GW (2017) The influence of monsoon climate on latewood growth of southwestern ponderosa pine. Forests 8(5): 140.
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Read PublicationThe North American Monsoon delivers warm season precipitation to much of the southwestern United States, yet the importance of this water source for forested ecosystems in the region is not well understood. While it is widely accepted that trees in southwestern forests use winter precipitation for earlywood production, the extent to which summer (monsoon season) precipitation supports latewood production is unclear. We used tree ring records, local climate data, and stable isotope analyses (δ<sup>18</sup>O) of water and cellulose to examine the importance of monsoon precipitation for latewood production in mature ponderosa pine (<i>Pinus ponderosa</i> Dougl.) in northern Arizona. Our analyses identified monsoon season vapor pressure deficit (VPD) and Palmer Drought Severity Index (PDSI) as significant effects on latewood growth, together explaining 39% of latewood ring width variation. Stem water and cellulose δ<sup>18</sup>O analyses suggest that monsoon precipitation was not directly used for latewood growth. Our findings suggest that mature ponderosa pines in this region utilize winter precipitation for growth throughout the entire year. The influence of monsoon precipitation on growth is indirect and mediated by its effect on atmospheric moisture stress (VPD). Together, summer VPD and antecedent soil moisture conditions have a strong influence on latewood growth.
Kim JS, Kug JS, Jeong SJ, Huntzinger DN, Michalak AM, Schwalm CR, Wei Y and Schaefer K (2017) Reduced North American terrestrial primary productivity linked to anomalous Arctic warming. Nature Geoscience 10: 572–576.
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Read PublicationWarming temperatures in the Northern Hemisphere have enhanced terrestrial productivity. Despite the warming trend, North America has experienced more frequent and more intense cold weather events during winters and springs. These events have been linked to anomalous Arctic warming since 1990, and may affect terrestrial processes. Here we analyse multiple observation data sets and numerical model simulations to evaluate links between Arctic temperatures and primary productivity in North America. We find that positive springtime temperature anomalies in the Arctic have led to negative anomalies in gross primary productivity over most of North America during the last three decades, which amount to a net productivity decline of 0.31 PgC yr−1across the continent. This decline is mainly explained by two factors: severe cold conditions in northern North America and lower precipitation in the South Central United States. In addition, United States crop-yield data reveal that during years experiencing anomalous warming in the Arctic, yields declined by approximately 1 to 4% on average, with individual states experiencing declines of up to 20%. We conclude that the strengthening of Arctic warming anomalies in the past decades has remotely reduced productivity over North America.
Klosterman S, Richardson AD (2017) Observing Spring and Fall Phenology in a Deciduous Forest with Aerial Drone Imagery. Sensors 17(12): 2852.
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Read PublicationPlant phenology is a sensitive indicator of the effects of global change on terrestrial ecosystems and controls the timing of key ecosystem functions including photosynthesis and transpiration. Aerial drone imagery and photogrammetric techniques promise to advance the study of phenology by enabling the creation of distortion-free orthomosaics of plant canopies at the landscape scale, but with branch-level image resolution. The main goal of this study is to determine the leaf life cycle events corresponding to phenological metrics derived from automated analyses based on color indices calculated from drone imagery. For an oak-dominated, temperate deciduous forest in the northeastern USA, we find that plant area index (PAI) correlates with a canopy greenness index during spring green-up, and a canopy redness index during autumn senescence. Additionally, greenness and redness metrics are significantly correlated with the timing of budburst and leaf expansion on individual trees in spring. However, we note that the specific color index for individual trees must be carefully chosen if new foliage in spring appears red, rather than green—which we observed for some oak trees. In autumn, both decreasing greenness and increasing redness correlate with leaf senescence. Maximum redness indicates the beginning of leaf fall, and the progression of leaf fall correlates with decreasing redness. We also find that cooler air temperature microclimates near a forest edge bordering a wetland advance the onset of senescence. These results demonstrate the use of drones for characterizing the organismic-level variability of phenology in a forested landscape and advance our understanding of which phenophase transitions correspond to color-based metrics derived from digital image analysis.
Koch BJ, Hungate BA, Price LB (2017) Food‐animal production and the spread of antibiotic resistance: The role of ecology. Frontiers in Ecology and the Environment 15(6): 309-318.
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Read PublicationAntibiotic‐resistant pathogens increasingly threaten human health. Widespread application of antibiotics to animal populations raised for food, including chickens, cattle, and pigs, selects for resistance and contributes to the evolution of those pathogens. Despite a half century of research establishing the mechanisms and pathways by which antibiotic‐resistant bacteria spread from food animals to people, scientists lack the appropriate data and models to estimate the public health burden of antibiotic‐resistant human infections attributable to antibiotic use in food‐animal production. Genomic technologies are enabling researchers to track the bidirectional transmissions of specific bacterial strains from livestock to people – and from people to livestock – that can amplify resistance traits. Concepts in ecology, which were developed to understand resource subsidies, metapopulations, and biological invasions, provide insight into the epidemiology of antibiotic resistance from genomic data. By applying ecological principles to highly resolved phylogenetic data, researchers can improve strategies for controlling antibiotic resistance.
Koirala S, Jung M, Reichstein M, Graaf IE, Camps‐Valls G, Ichii K, Papale D, Raduly B, Schwalm CR, Tramontana G and Carvalhais N (2017) Global distribution of groundwater-vegetation spatial covariation. Geophysical Research Letters 44(9): 4134–4142.
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Read PublicationGroundwater is an integral component of the water cycle, and it also influences the carbon cycle by supplying moisture to ecosystems. However, the extent and determinants of groundwater-vegetation interactions are poorly understood at the global scale. Using several high-resolution data products, we show that the spatial patterns of ecosystem gross primary productivity and groundwater table depth are correlated during at least one season in more than two thirds of the global vegetated area. Positive relationships, i.e., larger productivity under shallower groundwater table, predominate in moisture-limited dry to mesic conditions with herbaceous and shrub vegetation. Negative relationships, i.e., larger productivity under deeper groundwater, predominate in humid climates with forests, possibly indicating a drawdown of groundwater table due to substantial ecosystem water use. Interestingly, these opposite groundwater-vegetation interactions are primarily associated with differences in vegetation than with climate and surface characteristics. These findings put forth the first evidence, and a need for better representation, of extensive and non-negligible groundwater-vegetation interactions at the global scale.
Liu X-JA, Sun J, Mau RL, Finley BK, Compson ZG, van Gestel N, Brown JR, Schwartz E, Dijkstra P, Hungate BA (2017) Labile carbon inputs determines the direction and magnitude of the soil priming effect. Applied Soil Ecology 109: 7-13.
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Read PublicationLabile carbon (C) input to soil can accelerate or slow the decomposition of soil organic matter, a phenomenon called priming. However, priming is difficult to predict, making its relationship with C input elusive. To assess this relationship, we added <sup>13</sup>C-glucose at five levels (8 to 1606 μg C g<sup>−1</sup> week<sup>−1</sup>) to the soil from four different ecosystems for seven weeks. We observed a positive linear relationship between C input and priming in all soils: priming increased from negative or no priming at low C input to strong positive priming at high C input. However, the sensitivity of priming to C input varied among soils and between ways of expressing C input, and decreased with elevation. Positive substrate thresholds were detected in three soils (56 to 242 μg C g<sup>−1</sup> week<sup>−1</sup>), suggesting the minimum C input required to trigger positive priming. Carbon input expressed as a fraction of microbial biomass explained 16.5% less variation in priming than did C input expressed as a fraction of dry soil mass, indicating that priming is not strongly related to the size of the soil microbial biomass. We conclude that priming increases with the rate of labile C input, once that rate exceeds a threshold, but the magnitude of increase varies among soils. Further research on mechanisms causing the variation of priming sensitivity to increasing labile C input might help promote a quantitative understanding of how such phenomenon impacts soil C cycling, offering the potential to improve earth system models.
Liu XJ, van Groenigen KJ, Dijkstra P, Hungate BA (2017) Increased plant uptake of native soil nitrogen following fertilizer addition – not a priming effect?. Applied Soil Ecology 110: 105-110.
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Read PublicationFertilizer inputs affect plant uptake of native soil nitrogen (N), yet the underlying mechanisms remain elusive. To increase mechanistic insight into this phenomenon, we evaluated the effect of fertilizer addition on mineralization (in the absence of plants) and plant uptake of native soil N. We synthesized 43 isotope tracer (<sup>15</sup>N) studies and estimated the effects of fertilizer addition using <em>meta</em>-analysis. We found that organic fertilizer tended to reduce native soil N mineralization (−99 kg ha<sup>−1</sup> year<sup>−1</sup>; p = 0.09) while inorganic fertilizer tended to increase N priming (58 kg ha<sup>−1</sup> year<sup>−1</sup>; p = 0.17). In contrast, both organic and inorganic fertilizers significantly increased plant uptake of native soil N (179 and 107 kg ha<sup>−1</sup> year<sup>−1</sup>). Organic fertilizer had greater effect on plant uptake than on mineralization of native soil N (p < 0.001), but inorganic fertilizer had similar effects. Fertilizer effects on mineralization and plant uptake of native soil N were not influenced by study location (laboratory or field) and duration, soil texture, carbon and N content, and pH. Fertilizer addition variably affected native soil N mineralization but consistently increased plant uptake of native soil N. The positive effect of organic fertilizer on plant uptake of native soil N can not be explained by its negative effect on native soil N mineralization, suggesting that increased plant uptake of native soil N was caused mostly by plant-mediated mechanisms (e.g., increased root growth, rhizosphere N priming) rather than by soil microbe-mediated mechanisms.
Ma S, Jiang J, Huang Y, Shi Z, Wilson RM, Ricciuto D, Sebestyen SD, Hanson PJ, Luo Y (2017) Data‐constrained projections of methane fluxes in a northern Minnesota peatland in response to elevated CO2 and warming. Journal of Geophysical Research: Biogeosciences 122(11): 2841-2861.
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Read PublicationLarge uncertainties exist in predicting responses of wetland methane (CH4) fluxes to future climate change. However, sources of the uncertainty have not been clearly identified despite the fact that methane production and emission processes have been extensively explored. In this study, we took advantage of manual CH4 flux measurements under ambient environment from 2011 to 2014 at the Spruce and Peatland Responses Under Changing Environments (SPRUCE) experimental site and developed a data‐informed process‐based methane module. The module was incorporated into the Terrestrial ECOsystem (TECO) model before its parameters were constrained with multiple years of methane flux data for forecasting CH4 emission under five warming and two elevated CO2 treatments at SPRUCE. We found that 9°C warming treatments significantly increased methane emission by approximately 400%, and elevated CO2 treatments stimulated methane emission by 10.4%–23.6% in comparison with ambient conditions. The relative contribution of plant‐mediated transport to methane emission decreased from 96% at the control to 92% at the 9°C warming, largely to compensate for an increase in ebullition. The uncertainty in plant‐mediated transportation and ebullition increased with warming and contributed to the overall changes of emissions uncertainties. At the same time, our modeling results indicated a significant increase in the emitted CH4:CO2 ratio. This result, together with the larger warming potential of CH4, will lead to a strong positive feedback from terrestrial ecosystems to climate warming. The model‐data fusion approach used in this study enabled parameter estimation and uncertainty quantification for forecasting methane fluxes.
Mauritz M, Bracho R, Celis G, Hutchings J, Natali SM, Pegoraro E, Salmon VG, Schädel C, Webb EE, Schuur EAG (2017) Non-linear CO2 flux response to seven years of experimentally induced permafrost thaw. Global Change Biology.
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Read PublicationRapid Arctic warming is expected to increase global greenhouse gas concentrations as permafrost thaw exposes immense stores of frozen carbon (C) to microbial decomposition. Permafrost thaw also stimulates plant growth, which could offset C loss.Using data from seven years of experimental Air and Soil warming in moist acidic tundra, we show that Soil warming had a much stronger effect on CO2 flux than Air warming. Soil warming caused rapid permafrost thaw and increased ecosystem respiration (Reco), gross primary productivity (GPP), and net summer CO2 storage (NEE). Over seven years Reco, GPP, and NEE also increased in Control (i.e., ambient plots), but this change could be explained by slow thaw in Control areas. In the initial stages of thaw, Reco, GPP, and NEE increased linearly with thaw across all treatments, despite different rates of thaw. As thaw in Soil warming continued to increase linearly, ground surface subsidence created saturated micro-sites, and suppressed Reco, GPP, and NEE. However Reco and GPP remained high in areas with large Eriophorum vaginatum biomass. In general NEE increased with thaw, but was more strongly correlated with plant biomass than thaw, indicating that higher Reco in deeply thawed areas during summer months was balanced by GPP. Summer CO2 flux across treatments fit a single quadratic relationship that captured the functional response of CO<sub>2</sub> flux to thaw, water table depth, and plant biomass. These results demonstrate the importance of indirect thaw effects on CO2 flux: plant growth and water table dynamics. Non-summer Reco models estimated that the area was an annual CO2 source during all years of observation. Non-summer CO2 loss in warmer, more deeply thawed soils exceeded the increases in summer GPP, and thawed tundra was a net annual CO2 source.
McHugh, TA, Compson Z, van Gestel N, Hayer M, Ballard L, Haverty M, Hines, J, Irvine N, Krassner D, Lyons T, Musta EJ, Schiff M, Zint P, Schwartz E (2017) Climate controls prokaryotic community composition in desert soils of the southwestern United States. FEMS Microbiol Ecol.
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Read PublicationAridisols are the dominant soil type in drylands, which occupy one-third of Earth's terrestrial surface. We examined controls on biogeographical patterns of Aridisol prokaryotic (bacterial and archaeal) communities at a regional scale by comparing communities from 100 Aridisols throughout the southwestern United States using high-throughput sequencing of the 16S rRNA gene. We found that microbial communities differed among global biomes and deserts of the Southwest. Differences among biomes were driven by differences in taxonomic identities, whereas differences among deserts of the Southwest were driven by differences in relative sequence abundance. Desert communities were dominated by Actinobacteria, Proteobacteria and Crenarchaeota, supporting the notion of a core set of abundant taxa in desert soils. Our findings contrast with studies showing little taxonomic overlap at the OTU level (97% sequence similarity) across large spatial scales, as we found ∼90% of taxa in at least two of the three deserts. Geographic distance structured prokaryotic communities indirectly through the influence of climate and soil properties. Structural equation modeling suggests that climate exerts a stronger influence than soil properties in shaping the composition of Aridisol microbial communities, with annual heat moisture index (an aridity metric) being the strongest climate driver. Annual heat moisture index was associated with decreased microbial diversity and richness. If the Desert Southwest becomes hotter and drier as predicted, these findings suggest that prokaryotic diversity and richness in Aridisols will decline.
McLauchlan KK, Gerhart LM, Battles JJ, Craine JM, Elmore AJ, Higuera PE, Mack MC, McNeil BE, Nelson DM, Pederson N, Perakis SS (2017) Centennial-scale reductions in nitrogen availability in temperate forests of the United States. Scientific Reports 7: 7856.
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Read PublicationForests cover 30% of the terrestrial Earth surface and are a major component of the global carbon (C) cycle. Humans have doubled the amount of global reactive nitrogen (N), increasing deposition of N onto forests worldwide. However, other global changes—especially climate change and elevated atmospheric carbon dioxide concentrations—are increasing demand for N, the element limiting primary productivity in temperate forests, which could be reducing N availability. To determine the long-term, integrated effects of global changes on forest N cycling, we measured stable N isotopes in wood, a proxy for N supply relative to demand, on large spatial and temporal scales across the continental U.S.A. Here, we show that forest N availability has generally declined across much of the U.S. since at least 1850 C.E. with cool, wet forests demonstrating the greatest declines. Across sites, recent trajectories of N availability were independent of recent atmospheric N deposition rates, implying a minor role for modern N deposition on the trajectory of N status of North American forests. Our results demonstrate that current trends of global changes are likely to be consistent with forest oligotrophication into the foreseeable future, further constraining forest C fixation and potentially storage.
Melvin AM, Celis G, Johnsto, JF, McGuire AD, Gene, H, Schuur EAG, Rupp TS, Mack MC (2017) Fuel-reduction management alters plant composition, carbon and nitrogen pools, and soil thaw in Alaskan boreal forest. Ecological Applications 28(1):149-161.
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Read PublicationIncreasing wildfire activity in Alaska's boreal forests has led to greater fuel-reduction management. Management has been implemented to reduce wildfire spread, but the ecological impacts of these practices are poorly known. We quantified the effects of hand-thinning and shearblading on above- and belowground stand characteristics, plant species composition, carbon (C) and nitrogen (N) pools, and soil thaw across 19 sites dominated by black spruce (Picea mariana) in interior Alaska treated 2?12 years prior to sampling. The density of deciduous tree seedlings was significantly higher in shearbladed areas compared to unmanaged forest (6.4 vs. 0.1 stems/m2), and unmanaged stands exhibited the highest mean density of conifer seedlings and layers (1.4 stems/m2). Understory plant community composition was most similar between unmanaged and thinned stands. Shearblading resulted in a near complete loss of aboveground tree biomass C pools while thinning approximately halved the C pool size (1.2 kg C/m2 compared to 3.1 kg C/m2 in unmanaged forest). Significantly smaller soil organic layer (SOL) C and N pools were observed in shearbladed stands (3.2 kg C/m2 and 116.8 g N/m2) relative to thinned (6.0 kg C/m2 and 192.2 g N/m2) and unmanaged (5.9 kg C/m2 and 178.7 g N/m2) stands. No difference in C and N pool sizes in the uppermost 10 cm of mineral soil was observed among stand types. Total C stocks for measured pools was 2.6 kg C/m2 smaller in thinned stands and 5.8 kg C/m2 smaller in shearbladed stands when compared to unmanaged forest. Soil thaw depth averaged 13 cm deeper in thinned areas and 46 cm deeper in shearbladed areas relative to adjacent unmanaged stands, although variability was high across sites. Deeper soil thaw was linked to shallower SOL depth for unmanaged stands and both management types, however for any given SOL depth, thaw tended to be deeper in shearbladed areas compared to unmanaged forest. These findings indicate that fuel-reduction management alters plant community composition, C and N pools, and soil thaw depth, with consequences for ecosystem structure and function beyond those intended for fire management.
Morrissey EM, Mau RL, Schwartz E, McHugh TA, Dijkstra P, Koch BJ, Marks JC, and Hungate BA (2017) Bacterial carbon use plasticity, phylogenetic diversity and the priming of soil organic matter. ISME.
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Read PublicationMicroorganisms perform most decomposition on Earth, mediating carbon (C) loss from ecosystems, and thereby influencing climate. Yet, how variation in the identity and composition of microbial communities influences ecosystem C balance is far from clear. Using quantitative stable isotope probing of DNA, we show how individual bacterial taxa influence soil C cycling following the addition of labile C (glucose). Specifically, we show that increased decomposition of soil C in response to added glucose (positive priming) occurs as a phylogenetically diverse group of taxa, accounting for a large proportion of the bacterial community, shift toward additional soil C use for growth. Our findings suggest that many microbial taxa exhibit C use plasticity, as most taxa altered their use of glucose and soil organic matter depending upon environmental conditions. In contrast, bacteria that exhibit other responses to glucose (reduced growth or reliance on glucose for additional growth) clustered strongly by phylogeny. These results suggest that positive priming is likely the prototypical response of bacteria to sustained labile C addition, consistent with the widespread occurrence of the positive priming effect in nature.
O'Connor MI, Gonzalez A, Byrnes JEK, Cardinale BJ, Duffy JE, Gamfeldt L, Griffin JN, Hooper D, Hungate BA, Paquette A, Thompson PL, Dee LE, Dolan KL (2017) A general biodiversity-function relationship is mediated by trophic level. Oikos 126: 18-31.
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Read PublicationSpecies diversity affects the functioning of ecosystems, including the efficiency by which communities capture limited resources, produce biomass, recycle and retain biologically essential nutrients. These ecological functions ultimately support the ecosystem services upon which humanity depends. Despite hundreds of experimental tests of the effect of biodiversity on ecosystem function (BEF), it remains unclear whether diversity effects are sufficiently general that we can use a single relationship to quantitatively predict how changes in species richness alter an ecosystem function across trophic levels, ecosystems and ecological conditions. Our objective here is to determine whether a general relationship exists between biodiversity and standing biomass. We used hierarchical mixed effects models, based on a power function between species richness and biomass production (Y = a × S<sup>b</sup>), and a database of 374 published experiments to estimate the BEF relationship (the change in biomass with the addition of species), and its associated uncertainty, in the context of environmental factors. We found that the mean relationship (<em>b</em> = 0.26, 95% CI: 0.16, 0.37) characterized the vast majority of observations, was robust to differences in experimental design, and was independent of the range of species richness levels considered. However, the richness–biomass relationship varied by trophic level and among ecosystems; in aquatic systems <em>b</em> was nearly twice as large for consumers (herbivores and detritivores) compared to primary producers; in terrestrial ecosystems, <em>b</em> for detritivores was negative but depended on few studies. We estimated changes in biomass expected for a range of changes in species richness, highlighting that species loss has greater implications than species gains, skewing a distribution of biomass change relative to observed species richness change. When biomass provides a good proxy for processes that underpin ecosystem services, this relationship could be used as a step in modeling the production of ecosystem services and their dependence on biodiversity.
Prevey J, Vellend M, Ruger N, Hollister RD, Bjorkman AD, Myers-Smith IH, Elmendorf SC, Clark K, Cooper EJ, Elberling B, Fosaa AM, Henry GHR, Hoye TT, Jonsdottir IS, Klanderud K, Levesque E, Mauritz M, Molau U, Natali SM, Oberbauer SF, Panchen ZA, Post E, Rumpf SB, Schmidt NM, Schuur EAG, Semenchuk PR, Troxler T, Welker JM, Rixen C (2017) Greater temperature sensitivity of plant phenology at colder sites: implications for convergence across northern latitudes. Global Change Biology 23(7): 2660-2671.
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Read PublicationWarmer temperatures are accelerating the phenology of organisms around the world. Temperature sensitivity of phenology might be greater in colder, higher latitude sites than in warmer regions, in part because small changes in temperature constitute greater relative changes in thermal balance at colder sites. To test this hypothesis, we examined up to 20 years of phenology data for 47 tundra plant species at 18 high‐latitude sites along a climatic gradient. Across all species, the timing of leaf emergence and flowering was more sensitive to a given increase in summer temperature at colder than warmer high‐latitude locations. A similar pattern was seen over time for the flowering phenology of a widespread species, Cassiope tetragona. These are among the first results highlighting differential phenological responses of plants across a climatic gradient and suggest the possibility of convergence in flowering times and therefore an increase in gene flow across latitudes as the climate warms.
Price LB, Hungate BA, Koch BJ, Davis GS, Liu CM (2017) Colonizing opportunistic pathogens (COPs): The beasts in all of us. PloS Pathogens 13(8): e1006369.
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Read PublicationColonizing opportunistic pathogens (COPs) are microbes that asymptomatically colonize the human body and, when the conditions are right, can cause infections. Their ability to persist indefinitely and to be transmitted without detection gives COPs a unique epidemiology that warrants special consideration. There are examples of COPs among bacteria, fungi (e.g., Candida albicans), protozoa (e.g., Blastocystis), and viruses (e.g., Rhinovirus), but bacterial COPs are of particular relevance because of their major contribution to today’s antibiotic resistance crisis. The COPs include a long list of notorious bacteria that live double lives as passive stowaways and virulent foes. Some of the best-known COPs include Staphylococcus aureus, extraintestinal pathogenic Escherichia coli (ExPEC), Klebsiella pneumoniae, and Streptococcus pneumoniae (Table 1). Their capacity for benign coexistence with humans belies their alter egos that exact a heavy burden of human disease. For example, in the United States, ExPEC bloodstream infections kill as many as 40,000 people annually, but, ExPEC are also benign colonizers in the gastrointestinal tract. Host factors, including age, sex, health status, anatomy, and behavior, all play profound roles in infection susceptibility and severity. In particular, immunocompromised individuals are at excess risk for infections caused by diverse bacteria, including COPs and even commensals. Yet, health status is not the sole determinant of infection by COPs. For example, healthy women more frequently suffer from urinary tract infections than men because of anatomical differences, including shorter urethrae. Likewise, healthy children more commonly suffer from acute otitis media than adults due to their shorter, flatter eustachian tubes.
Robinson NP, Allred BW, Jones MO, Moreno A, Kimball JS, Naugle DE, Erickson TA, Richardson AD (2017) A Dynamic Landsat Derived Normalized Difference Vegetation Index (NDVI) product for the conterminous United States. Remote Sensing 9(8): 863.
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Read PublicationSatellite derived vegetation indices (VIs) are broadly used in ecological research, ecosystem modeling, and land surface monitoring. The Normalized Difference Vegetation Index (NDVI), perhaps the most utilized VI, has countless applications across ecology, forestry, agriculture, wildlife, biodiversity, and other disciplines. Calculating satellite derived NDVI is not always straight-forward, however, as satellite remote sensing datasets are inherently noisy due to cloud and atmospheric contamination, data processing failures, and instrument malfunction. Readily available NDVI products that account for these complexities are generally at coarse resolution; high resolution NDVI datasets are not conveniently accessible and developing them often presents numerous technical and methodological challenges. We address this deficiency by producing a Landsat derived, high resolution (30 m), long-term (30+ years) NDVI dataset for the conterminous United States. We use Google Earth Engine, a planetary-scale cloud-based geospatial analysis platform, for processing the Landsat data and distributing the final dataset. We use a climatology driven approach to fill missing data and validate the dataset with established remote sensing products at multiple scales. We provide access to the composites through a simple web application, allowing users to customize key parameters appropriate for their application, question, and region of interest.
Schwalm CR, Anderegg WRL, Michalak AM, Fisher JB, Biondi F, Koch G, Litvak M, Ogle K, Shaw JD, Wolf A, Huntzinger DN, Schaefer K, Cook R, Wei Y, Fang Y, Hayes D, Huang M, Jain A, Tian H (2017) Global patterns of drought recovery. Nature 548: 202-205.
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Read PublicationDrought, a recurring phenomenon with major impacts on both human and natural systems, is the most widespread climatic extreme that negatively affects the land carbon sink. Although twentieth-century trends in drought regimes are ambiguous across many regions more frequent and severe droughts are expected in the twenty-first century. Recovery time—how long an ecosystem requires to revert to its pre-drought functional state—is a critical metric of drought impact. Yet the factors influencing drought recovery and its spatiotemporal patterns at the global scale are largely unknown. Here we analyse three independent datasets of gross primary productivity and show that, across diverse ecosystems, drought recovery times are strongly associated with climate and carbon cycle dynamics, with biodiversity and CO2 fertilization as secondary factors. Our analysis also provides two key insights into the spatiotemporal patterns of drought recovery time: first, that recovery is longest in the tropics and high northern latitudes (both vulnerable areas of Earth’s climate system) and second, that drought impacts (assessed using the area of ecosystems actively recovering and time to recovery) have increased over the twentieth century. If droughts become more frequent, as expected, the time between droughts may become shorter than drought recovery time, leading to permanently damaged ecosystems and widespread degradation of the land carbon sink.
Strauss J, Schirrmeister L, Grosse G, Fortier D, Hugelius G, Knoblauch C, Romanovsky V, Schädel C, Schneider von Deimling T, Schuur EAG, Shmelev D, Ulrich M, Veremeeva A (2017) Deep Yedoma permafrost: A synthesis of depositional characteristics and carbon vulnerability. Earth-Science Reviews 13: 83-95.
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Read PublicationPermafrost is a distinct feature of the terrestrial Arctic and is vulnerable to climate warming. Permafrost degrades in different ways, including deepening of a seasonally unfrozen surface and localized but rapid development of deep thaw features. Pleistocene ice-rich permafrost with syngenetic ice-wedges, termed Yedoma deposits, are widespread in Siberia, Alaska, and Yukon, Canada and may be especially prone to rapid-thaw processes. Freeze-locked organic matter in such deposits can be re-mobilized on short time-scales and contribute to a carbon-cycle climate feedback. Here we synthesize the characteristics and vulnerability of Yedoma deposits by synthesizing studies on the Yedoma origin and the associated organic carbon pool. We suggest that Yedoma deposits accumulated under periglacial weathering, transport, and deposition dynamics in non-glaciated regions during the late Pleistocene until the beginning of late glacial warming. The deposits formed due to a combination of aeolian, colluvial, nival, and alluvial deposition and simultaneous ground ice accumulation. We found up to 130 gigatons organic carbon in Yedoma, parts of which are well-preserved and available for fast decomposition after thaw. Based on incubation experiments, up to 10% of the Yedoma carbon is considered especially decomposable and may be released upon thaw. The substantial amount of ground ice in Yedoma makes it highly vulnerable to disturbances such as thermokarst and thermo-erosion processes. Mobilization of permafrost carbon is expected to increase under future climate warming. Our synthesis results underline the need of accounting for Yedoma carbon stocks in next generation Earth-System-Models for a more complete representation of the permafrost-carbon feedback.
Taylor MA, Hendy IL, Chappaz A (2017) Assessing oxygen depletion in the Northeastern Pacific Ocean during the last deglaciation using I/Ca ratios from multiple benthic foraminiferal species. Paleoceanography 32(8): 746–762.
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Read PublicationPaleo-redox proxies are crucial for reconstructing past bottom water oxygen concentration changes brought about by ocean circulation and marine productivity shifts in response to climate forcing. Carbonate I/Ca ratios of multiple benthic foraminifera species from Ocean Drilling Program Hole 1017E—a core drilled within the Californian oxygen minimum zone (OMZ), on the continental slope—are employed to reexamine the transition from the well-oxygenated last glacial into poorly oxygenated modern conditions. The redox and export productivity history of this site is constrained by numerous proxies used to assess sensitivity of I/Ca ratios of benthic foraminifera to changes in bottom and pore water O2 concentrations. Reconstructed iodate (IO3−) availability is from the I/Ca ratio of epifaunal (Cibicidoides sp.), shallow infaunal (Uvigerina peregrina), and deep infaunal (Bolivina spissa) foraminifera. The reconstructed IO3− availability profile is used to determine the contribution of bottom water O2 relative to oxidant demand on pore water O2 concentrations. These results suggest that high export productivity on the California Margin drove low pore water O2 concentrations during the Bølling. In contrast, low bottom water O2 concentrations at 950 m water depth only contributed to reduced sediments during the Allerød. Increased contribution of modified North Pacific Intermediate Water to the California Current System ventilated the California OMZ during the late glacial and the Younger Dryas such that water overlying the site was oxygenated. These results highlight the promising potential of this new proxy for understanding the relative influence of bottom water O2concentration and pore water oxidant demand on OMZs.
Turetsky MR, Baltzer JL, Johnstone JF, Mack MC, McCann K, Schuur EAG, (2017) Losing legacies, ecological release, and transient responses: Key challenges for the future of northern ecosystem science. Ecosystems 20(1): 23-30.
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Read PublicationNorthern ecosystem processes play out across scales that are rare elsewhere on contemporary earth: large ranging predator–prey systems are still operational, invasive species are rare, and large-scale natural disturbances occur extensively. Disturbances in the far north affect huge areas of land and are difficult to control or manage. Historically, disturbance patterns and processes ranging across a number of spatio-temporal scales have played an important role in the resilience of northern ecosystems. However, due to interactions with a warming climate, these disturbances are now erasing key legacies of the last millennia of ecosystem processes. Building on the concepts of legacies and cross-scale interactions, we highlight several general conceptual issues that represent key challenges for the future of northern ecosystem science, but that also have relevance to other biomes.
Van Groenigen JW, Van Kessel C, Hungate BA, Oenema O, Powlson DS, Van Groenigen KJ (2017) Response to the Letter to the Editor regarding our viewpoint “Sequestering soil organic carbon: A nitrogen dilemma”. Environmental Science & Technology 51(20): 11503-11504.
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Read PublicationWe disagree with the statement by Soussana et al. that the 4p1000 goal is already sufficiently spatially diversified because it is related to the local soil organic C (SOC) stock. This implies that soils with a large SOC stock will normally have a larger nitrogen (N) (and phosphorus, P) surplus than those containing less SOC. We fail to see the rationale for their statement in two ways. First, at the global scale, many soils with a large SOC stock will be (extensively) grazed grasslands, which typically have small inputs of N4 and P inputs and small surpluses. In contrast, many intensively managed arable soils, which typically have lower SOC stocks, have large inputs of N and P leading to large surpluses. Second, in general, soils with a low SOC stock (e.g., old arable soils, degraded lands, mine wastes) have greater potential for increasing SOC than soils with high SOC stocks. Focusing C sequestration efforts on these soils would seem advantageous, both for climate change mitigation and for improving soil quality.
van Groenigen KJ, Osenberg CW, Terrer C, Carrillo Y, Dijkstra F, Heath J, Nie M, Pendall E, Phillips RP, Hungate BA (2017) Faster turnover of new soil carbon inputs under increased atmospheric CO2. Global Change Biology.
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Read PublicationRising levels of atmospheric CO<sub>2</sub> frequently stimulate plant inputs to soil, but the consequences of these changes for soil carbon (C) dynamics are poorly understood. Plant-derived inputs can accumulate in the soil and become part of the soil C pool (“new soil C”), or accelerate losses of pre-existing (“old”) soil C. The dynamics of the new and old pools will likely differ and alter the long-term fate of soil C, but these separate pools, which can be distinguished through isotopic labeling, have not been considered in past syntheses. Using meta-analysis, we found that while elevated CO<sub>2</sub> (ranging from 550 to 800 parts per million by volume) stimulates the accumulation of new soil C in the short term (< 1 year), these effects do not persist in the longer term (1 - 4 years). Elevated CO<sub>2</sub> does not affect the decomposition or the size of the old soil C pool over either temporal scale. Our results are inconsistent with predictions of conventional soil C models and suggest that elevated CO<sub>2</sub> might increase turnover rates of new soil C. Because increased turnover rates of new soil C limit the potential for additional soil C sequestration, the capacity of land ecosystems to slow the rise in atmospheric CO<sub>2</sub> concentrations may be smaller than previously assumed.
Walker X, Frey MD, Conway AJ, Jean M, Johnstone JF (2017) Impacts of fire on non-native plant recruitment in black spruce forests of interior Alaska. PLOS ONE 12(2): e0171599.
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Read PublicationClimate change is expected to increase the extent and severity of wildfires throughout the boreal forest. Historically, black spruce (Picea mariana (Mill.) B.S.P.) forests in interior Alaska have been relatively free of non-native species, but the compounding effects of climate change and an altered fire regime could facilitate the expansion of non-native plants. We tested the effects of wildfire on non-native plant colonization by conducting a seeding experiment of non-native plants on different substrate types in a burned black spruce forest, and surveying for non-native plants in recently burned and mature black spruce forests. We found few non-native plants in burned or mature forests, despite their high roadside presence, although invasion of some burned sites by dandelion (Taraxacum officinale) indicated the potential for non-native plants to move into burned forest. Experimental germination rates were significantly higher on mineral soil compared to organic soil, indicating that severe fires that combust much of the organic layer could increase the potential for non-native plant colonization. We conclude that fire disturbances that remove the organic layer could facilitate the invasion of non-native plants providing there is a viable seed source and dispersal vector.
Wilson CH, Caughlin TT, Rifai SW, Boughton EH, Mack MC, Flory SL (2017) Multi-decadal time series of remotely sensed vegetation improves prediction of soil carbon in a subtropical grassland. Ecological Applications 27(5): 1646-1656.
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Read PublicationSoil carbon sequestration in agroecosystems could play a key role in climate change mitigation but will require accurate predictions of soil organic carbon (SOC) stocks over spatial scales relevant to land management. Spatial variation in underlying drivers of SOC, such as plant productivity and soil mineralogy, complicates these predictions. Recent advances in the availability of remotely sensed data make it practical to generate multidecadal time series of vegetation indices with high spatial resolution and coverage. However, the utility of such data largely is unknown, only having been tested with shorter (e.g., 1–2 yr) data summaries. Across a 2,000 ha subtropical grassland, we found that a long time series (28 yr) of a vegetation index (Enhanced Vegetation Index; EVI) derived from the Landsat 5 satellite significantly enhanced prediction of spatially varying SOC pools, while a short summary (2 yr) was an ineffective predictor. EVI was the best predictor for surface SOC (0–5 cm depth) and total measured SOC stocks (0–15 cm). The optimum models for SOC in the upper soil layer combined EVI records with elevation and calcium concentration, while deeper SOC was more strongly associated with calcium availability. We demonstrate how data from the open access Landsat archive can predict SOC stocks, a key ecosystem metric, and illustrate the rich variety of analytical approaches that can be applied to long time series of remotely sensed greenness. Overall, our results showed that SOC pools were closely coupled to EVI in this ecosystem, demonstrating that maintenance of higher average green leaf area is correlated with higher SOC. The strong associations of vegetation greenness and calcium concentration with SOC suggest that the ability to sequester additional SOC likely will rely on strategic management of pasture vegetation and soil fertility.
Yan Y, Zhou X, Jiang L, Luo Y (2017) Effects of carbon turnover time on terrestrial ecosystem carbon storage. Biogeosciences 14(23): 5441-5454.
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Read PublicationCarbon (C) turnover time is a key factor in determining C storage capacity in various plant and soil pools as well as terrestrial C sink in a changing climate. However, the effects of C turnover time on ecosystem C storage have not been well explored. In this study, we compared mean C turnover times (MTTs) of ecosystem and soil, examined their variability to climate, and then quantified the spatial variation in ecosystem C storage over time from changes in C turnover time and/or net primary production (NPP). Our results showed that mean ecosystem MTT based on gross primary production (GPP; MTTEC_GPP = Cpool/GPP, 25.0 ± 2.7 years) was shorter than soil MTT (MTTsoil = Csoil/NPP, 35.5 ± 1.2 years) and NPP-based ecosystem MTT (MTTEC_NPP = Cpool/NPP, 50.8 ± 3 years; Cpool and Csoil referred to ecosystem or soil C storage, respectively). On the biome scale, temperature is the best predictor for MTTEC (R2 = 0.77, p < 0.001) and MTTsoil (R2 = 0.68, p < 0.001), while the inclusion of precipitation in the model did not improve the performance of MTTEC (R2 = 0.76, p < 0.001). Ecosystem MTT decreased by approximately 4 years from 1901 to 2011 when only temperature was considered, resulting in a large C release from terrestrial ecosystems. The resultant terrestrial C release caused by the decrease in MTT only accounted for about 13.5 % of that due to the change in NPP uptake (159.3 ± 1.45 vs. 1215.4 ± 11.0 Pg C). However, the larger uncertainties in the spatial variation of MTT than temporal changes could lead to a greater impact on ecosystem C storage, which deserves further study in the future.
Zhang X, Hutchings JA, Bianchi TS, Liu Y, Arellano AR, Schuur EAG (2017) Importance of lateral flux and its percolation depth on organic carbon export in Arctic tundra soil: Implications from a soil leaching experiment. Biogeosciences 122(4): 796-810.
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Read PublicationTemperature rise in the Arctic is causing deepening of active layers and resulting in the mobilization of deep permafrost dissolved organic matter (DOM). However, the mechanisms of DOM mobilization from Arctic soils, especially upper soil horizons which are drained most frequently through a year, are poorly understood. Here we conducted a short-term leaching experiment on surface and deep organic active layer soils, from the Yukon River basin, to examine the effects of DOM transport on bulk and molecular characteristics. Our data showed a net release of DOM from surface soils equal to an average of 5% of soil carbon. Conversely, deep soils percolated with surface leachates retained up to 27% of bulk DOM while releasing fluorescent components (up to 107%), indicating selective release of aromatic components (e.g., lignin and tannin), while retaining nonchromophoric components, as supported by spectrofluorometric and ultrahigh-resolution mass spectroscopic techniques. Our findings highlight the importance of the lateral flux of DOM on ecosystem carbon balance as well as processing of DOM transport through organic active layer soils en route to rivers and streams. This work also suggests the potential role of leachate export as an important mechanism of C losses from Arctic soils, in comparison with the more traditional pathway from soil to atmosphere in a warming Arctic.
Zhang X, Jayavelu S, Liu L, Friedl M, Henebry G, Liu Y, Schaaf CB, Richardson AD, Gray J (2017) Evaluation of land surface phenology from VIIRS data using time series of PhenoCam imagery. Agricultural And Forest Meteorology 256-257, 137-149.
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Read PublicationLand surface phenology (LSP) has been widely retrieved from time series of various satellite instruments in order to monitor climate change and ecosystem dynamics. However, any evaluation of the quality of LSP data sets is quite challenging because the in situ observations on a limited number of individual trees, shrubs, or other plants are rarely representative of the landscape sampled in a single satellite pixel. Moreover, vegetation indices detecting biophysical features of vegetation seasonality are different from (but related to) the specific plant life history stages observed by humans at ground level. This study is the first comprehensive evaluation of the LSP product derived from Visible Infrared Imaging Radiometer Suite (VIIRS) data using both MODIS LSP products and observations from the PhenoCam network across the Contiguous United States during 2013 and 2014. PhenoCam observes vegetation canopy over a landscape at very high frequency, providing nearly continuous canopy status and enabling the estimate of discrete phenophase using vegetation indices that are conceptually similar to satellite data. Six phenological dates (greenup onset, mid-greenup phase, maturity onset, senescence onset, mid-senescence phase, and dormancy onset) were retrieved separately from daily VIIRS NDVI (normalized difference vegetative index) and EVI2 (two-band enhanced vegetation index) time series. Similarly, the six phenological dates were also extracted from the 30-min time series of PhenoCam data using GCC (green chromatic coordinate) and VCI (vegetation contrast index) separately. Phenological dates derived from VIIRS NDVI and EVI2 and PhenoCam GCC and VCI were generally comparable for the vegetation greenup phase, but differed considerably for the senescence phase. Although all indices captured green leaf development effectively, performance discrepancies arose due to their ability to track the mixture of senescing leaf colors. PhenoCam GCC and VCI phenological observations were in better agreement with the phenological dates from VIIRS EVI2 than from VIIRS NDVI. Further, the VIIRS EVI2 phenological metrics were more similar to those from PhenoCam VCI than from PhenoCam GCC time series. Overall, the average absolute difference between the VIIRS EVI2 and PhenoCam VCI phenological dates was 7–11 days in the greenup phase and 10–13 days in the senescence phase. The difference was smaller in forests, followed by grasslands and croplands, and then savannas. Finally, the phenological dates derived from VIIRS EVI2 were compared with MODIS detections, which showed a good agreement with an average absolute difference less than a week except for the senescence onset. These results for the first time demonstrate the upper boundary of uncertainty in VIIRS LSP detections and the continuity to MODIS LSP product.
Zhou S, Yu B, Schwalm CR, Ciais P, Zhang Y, Fisher JB, Michalak AM, Wang W, Poulter B, Huntzinger DN, Niu S, Mao J, Jain A, Ricciuto DM, Shi X, Ito A, Wei Y, Huang Y, Wang G (2017) Response of water use efficiency to global environmental change based on output from terrestrial biosphere models. Global Biogeochemical Cycles 31(11): 1639-1655.
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Read PublicationWater use efficiency (WUE), defined as the ratio of gross primary productivity and evapotranspiration at the ecosystem scale, is a critical variable linking the carbon and water cycles. Incorporating a dependency on vapor pressure deficit, apparent underlying WUE (uWUE) provides a better indicator of how terrestrial ecosystems respond to environmental changes than other WUE formulations. Here we used 20th century simulations from four terrestrial biosphere models to develop a novel variance decomposition method. With this method, we attributed variations in apparent uWUE to both the trend and interannual variation of environmental drivers. The secular increase in atmospheric CO2 explained a clear majority of total variation (66 ± 32%: mean ± one standard deviation), followed by positive trends in nitrogen deposition and climate, as well as a negative trend in land use change. In contrast, interannual variation was mostly driven by interannual climate variability. To analyze the mechanism of the CO2 effect, we partitioned the apparent uWUE into the transpiration ratio (transpiration over evapotranspiration) and potential uWUE. The relative increase in potential uWUE parallels that of CO2, but this direct CO2 effect was offset by 20 ± 4% by changes in ecosystem structure, that is, leaf area index for different vegetation types. However, the decrease in transpiration due to stomatal closure with rising CO2 was reduced by 84% by an increase in leaf area index, resulting in small changes in the transpiration ratio. CO2 concentration thus plays a dominant role in driving apparent uWUE variations over time, but its role differs for the two constituent components: potential uWUE and transpiration.
2016
Abbott BW, Jones JBJ, Schuur EAG, Chapin FSI, Bowden WB, Bret-Harte MS, Epstein HE, Flannigan MD, Harms TK, Hollingsworth TN, Mack MC, McGuire AD, Natali SM, Rocha AV, Tank SE, Turetsky MR, Vonk JE, Wickland KP, Aiken GR, Alexander HD, Amon RMW, Bensoter BW, Bergeron Y, Bishop K, Blarquez O, Bond-Lamberty B, Breen AL, Buffam I, Cai Y, Carcaillet C, Carey SK, Chen JM, Chen HYH, Christensen TR, Cooper LW, Cornelissen JHC, de Groot WJ, DeLuca TH, Dorrepaal E, Fetcher N, Finlay JC, Forbes BC, French NHF, Gauthier S, Girardin MP, Goetz SJ, Goldammer JG, Gouch L, Grogan P, Guo L, Higuera PE, Hinzman L, Hu FS, Hugelius G, Jafarov EE, Jandt R, Johnstone JF, Karlsson J, Kasischke ES, Kattner G, Kelly R, Keuper F, Kling GW, Kortelainen P, Kouki J, Kuhry P, Laudon H, Laurion I, Macdonald RW, Mann PJ, Martikainen PJ, McClelland JW, Molau U, Oberbauer SF, Olefeldt D, Paré D, Parisien M-A, Payette S, Peng C, Pokrovksy OS, Rastetter EB, Raymond PA, Raynolds MK, Rein G, Reynolds JF, Robard M, Rogers BM, Schädel C, Schaefer K, Schmidt IK, Shvidenko A, Sky J, Spencer RGM, Starr G, Striegl RG, Teisserenc R, Tranvik LJ, Virtanen T, Welker JM, Zimov S (2016) Biomass offsets little or none of permafrost carbon release from soils, streams, and wildfire: an expert assessment. Environmental Research Letters (11) 3, 034014.
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Read PublicationAs 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.
Alexander HD, Mack MC (2016) A Canopy Shift in Interior Alaskan Boreal Forests: Consequences for Above- and Belowground Carbon and Nitrogen Pools during Post-fire Succession. Ecosystems 19 (1): 98–114.
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Read PublicationGlobal change models predict that high-latitude boreal forests will become increasingly susceptible to fire activity as climate warms, possibly causing a positive feedback to warming through fire-driven emissions of CO<sub>2</sub> into the atmosphere. However, fire-climate feedbacks depend on forest regrowth and carbon (C) accumulation over the post-fire successional interval, which is influenced by nitrogen (N) availability. To improve our understanding of post-fire C and N accumulation patterns in boreal forests, we evaluated above- and belowground C and N pools within 70 stands throughout interior Alaska, a region predicted to undergo a shift in canopy dominance as fire severity increases. Stands represented gradients in age and successional trajectory, from black spruce (<em class="EmphasisTypeItalic ">Picea mariana</em>) self-replacement to species replacement by deciduous species of trembling aspen (<em class="EmphasisTypeItalic ">Populus tremuloides</em>) and Alaska paper birch (<em class="EmphasisTypeItalic ">Betula neoalaskana</em>). Stands undergoing deciduous trajectories stored proportionally more of their C and N in aboveground stemwood and had 5–7 times faster rates of aboveground net primary productivity of trees compared to stands undergoing a black spruce trajectory, which stored more of their C and N in the soil organic layer (SOL), a thick layer of mostly undecomposed mosses. Thus, as successional trajectories shift, total C and N pool sizes will remain relatively unchanged, but there will be a trade-off in pool location and a potential increase in C and N longevity due to decreased flammability and decomposition rates of deciduous stemwood. Despite often warmer, drier conditions in deciduous compared to black spruce stands, deciduous stemwood has a C:N around 10 times higher than the black spruce SOL and often remains standing for many years with reduced exposure to fungal decomposers. Thus, a fire-driven shift in successional trajectories could cause a negative feedback to climate warming because of increased pool longevity in deciduous trajectories.
Ambrose AR, Baxter WL, Wong CS, Burgess SSO, Williams CB, Næsborg RR, Koch GW, Dawson TE (2016) Hydraulic constraints modify optimal photosynthetic profiles in giant sequoia trees. Oecologia 182 (3): 713-730.
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Read PublicationOptimality theory states that whole-tree carbon gain is maximized when leaf N and photosynthetic capacity profiles are distributed along vertical light gradients such that the marginal gain of nitrogen investment is identical among leaves. However, observed photosynthetic N gradients in trees do not follow this prediction, and the causes for this apparent discrepancy remain uncertain. Our objective was to evaluate how hydraulic limitations potentially modify crown-level optimization in Sequoiadendron giganteum (giant sequoia) trees up to 90 m tall. Leaf water potential (Ψ l ) and branch sap flow closely followed diurnal patterns of solar radiation throughout each tree crown. Minimum leaf water potential correlated negatively with height above ground, while leaf mass per area (LMA), shoot mass per area (SMA), leaf nitrogen content (%N), and bulk leaf stable carbon isotope ratios (δ13C) correlated positively with height. We found no significant vertical trends in maximum leaf photosynthesis (A), stomatal conductance (g s), and intrinsic water-use efficiency (A/g s), nor in branch-averaged transpiration (<em>E</em>L), stomatal conductance (<i><span style="font-weight: 400;">G</span></i><span style="font-weight: 400;">S</span>), and hydraulic conductance (<em>K</em>L). Adjustments in hydraulic architecture appear to partially compensate for increasing hydraulic limitations with height in giant sequoia, allowing them to sustain global maximum summer water use rates exceeding 2000 kg day−1. However, we found that leaf N and photosynthetic capacity do not follow the vertical light gradient, supporting the hypothesis that increasing limitations on water transport capacity with height modify photosynthetic optimization in tall trees.
Bernal B, McKinley DC, Hungate BA, White PM, Mozdzer TJ, Megonigal JP (2016) Limits to soil carbon stability; Deep, ancient soil carbon decomposition stimulated by new labile organic inputs. Soil Biology and Biochemistry 98: 85-94.
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Read PublicationCarbon (C) buried deep in soil (below 1 m) is often hundreds to thousands of years old, though the stability and sensitivity of this deep C to environmental change are not well understood. We examined the C dynamics in three soil horizons and their responses to changes in substrate availability in a coarse-textured sandy spodosol (0.0–0.1, 1.0–1.3, and 2.7–3.0 m deep). Substrate additions were intended to mimic an increase in root exudates and available inorganic nitrogen (N) that would follow an increase of belowground biomass at depth, as previously found in a long-term CO<sub>2</sub> enrichment experiment at this site. We incubated these soils for 60 days with glucose, alanine, and leaf litter, crossed with an inorganic N amendment equivalent to three times ambient concentrations. The organic substrates were isotopically labeled (<sup>13</sup>C), allowing us to determine the source of mineralized C and assess the priming effect. Enzyme activity increased as much as 13 times in the two deeper horizons (1.0–1.3, and 2.7–3.0 m) after the addition of the organic substrates, even though the deepest horizon had microbial biomass and microbial phospholipid fatty acids below the level of detection before the experiment. The deepest horizon (2.7–3.0 m) yielded the largest priming response under alanine, indicating that microorganisms in these soil horizons can become active in response to input of organic substrates. Inorganic N amendments significantly decreased the priming effect, suggesting that decomposition may not be N limited. However, alanine (organic N) yielded the highest priming effect at every soil depth, indicating the importance of differentiating effect of organic and inorganic N on decomposition. Distinct priming effects with depth suggest that portions of the soil profile can respond differently to organic inputs. Our findings indicate that the deep soil C pools might be more vulnerable to environmental or anthropogenic change than previously thought, potentially influencing net CO<sub>2</sub> exchange estimates between the land and the atmosphere.
Bracho R, Natali S, Pegoraro E, Crummer KG, Schädel C, Celis G, Hale L, Wu L, Yin H, Tiedje JM, Konstantinidis KT, Luo Y, Zhou J, Schuur EAG (2016) Temperature sensitivity of organic matter decomposition of permafrost-region soils during laboratory incubations. Soil Biology and Biochemistry 97, 1-14.
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Read PublicationPermafrost soils contain more than 1300 Pg of carbon (C), twice the amount of C in the atmosphere. Temperatures in higher latitudes are increasing, inducing permafrost thaw and subsequent microbial decomposition of previously frozen C, which will most likely feed back to climate warming through release of the greenhouse gases CO2 and CH4. Understanding the temperature sensitivity (Q10) and dynamics of soil organic matter (SOM) decomposition under warming is essential to predict the future state of the climate system. Alaskan tundra soils from the discontinuous permafrost zone were exposed to in situ experimental warming for two consecutive winters, increasing soil temperature by 2.3 °C down to 40 cm in the soil profile. Soils obtained at three depths (0–15, 15–25 and 45–55 cm) from the experimental warming site were incubated under aerobic conditions at 15 °C and 25 °C over 365 days in the laboratory. Carbon fluxes were measured periodically and dynamics of SOM decomposition, C pool sizes, and decay rates were estimated. Q10 was estimated using both a short-term temperature manipulation (Q10-ST) performed at 14, 100 and 280 days of incubation and via the equal C method (Q10-EC, ratio of time taken for a soil to respire a given amount of C), calculated continuously. At the same time points, functional diversities of the soil microbial communities were monitored for all incubation samples using a microbial functional gene array, GeoChip 5.0. Each array contains over 80,000 probes targeting microbial functional genes involved in biogeochemical cycling of major nutrients, remediation strategies, pathogenicity and other important environmental functions. Of these, over 20,000 probes target genes involved in the degradation of varying C substrates and can be used to quantify the relative gene abundances and functional gene diversities related to soil organic matter turnover. The slow decomposing C pool (CS), which represented close to 95% of total C in the top 25 cm soils, had a higher Q10 than the fast decomposing C pool (CF) and also dominated the total amount of C released by the end of the incubation. Overall, CS had temperature sensitivities of Q10-ST = 2.55 ± 0.03 and Q10-EC = 2.19 ± 0.13, while the CF had a temperature sensitivity of Q10-EC = 1.16 ± 0.30. In contrast to the 15 °C incubations, the 25 °C microbial communities showed reduced diversities of C-degradation functional genes in the early stage of the incubations. However, as the incubations continued the 25 °C communities more closely paralleled the 15 °C communities with respect to the detection of microbial genes utilized in the degradation of labile to recalcitrant C substrates. Two winter seasons of experimental warming did not affect the dynamics and temperature sensitivity of SOM decomposition or the microbial C-degradation genes during incubation. However, under the projected sustained warming attributable to climate change, we might expect increased contribution of CS to organic matter decomposition. Because of the higher Q10 and the large pool size of CS, increased soil organic matter release under warmer temperatures will contribute towards accelerating climate change.
Buelow HN, Winter AS, Van Horn DJ, Barrett JE, Gooseff MN, Schwartz E, Takacs-Vesbach CD (2016) Microbial Community Responses to Increased Water and Organic Matter in the Arid Soils of the McMurdo Dry Valleys, Antarctica. Frontiers in Microbiology 7 (1040).
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Read Publication<span id="yui_3_14_1_1_1470075850807_2152" class="foldable-text" data-reactid="143"><span class="text-with-line-breaks">The soils of the McMurdo Dry Valleys, Antarctica are an extreme polar desert, inhabited exclusively by microscopic taxa. This region is on the threshold of anticipated climate change, with glacial melt, permafrost thaw, and the melting of massive buried ice increasing liquid water availability and mobilizing soil nutrients. Experimental water and organic matter (OM) amendments were applied to investigate how these climate change effects may impact the soil communities. To identify active taxa and their functions, total community RNA transcripts were sequenced and annotated, and amended soils were compared with unamended control soils using differential abundance and expression analyses. Overall, taxonomic diversity declined with amendments of water and organic matter. The domain Bacteria increased with both amendments while Eukaryota declined from 38% of all taxa in control soils to 8% and 11% in water and OM amended soils, respectively. Among bacterial phyla, Actinobacteria (59%) dominated water-amended soils and Firmicutes (45%) dominated OM amended soils. Three bacterial phyla (Actinobacteria, Proteobacteria, and Firmicutes) were primarily responsible for the observed positive functional responses, while eukaryotic taxa experienced the majority (27 of 34) of significant transcript losses. These results indicated that as climate changes in this region, a replacement of endemic taxa adapted to dry, oligotrophic conditions by generalist, copiotrophic taxa is likely.</span></span>
Butterfield BJ, Camhi AL, Rubin RL, and Schwalm CR (2016) Tradeoffs and Compatibilities Among Ecosystem Services: Biological, Physical and Economic Drivers of Multifunctionality. Advances in Ecological Research 54: 207-243.
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Read Publication<span id="yui_3_14_1_1_1461966615416_1099" class="foldable-text" data-reactid=".5aqlq823uo.1.0.0.0.4.1.0"><span id="yui_3_14_1_1_1461966615416_1253" class="text-with-line-breaks" data-reactid=".5aqlq823uo.1.0.0.0.4.1.0.0"><span id="yui_3_14_1_1_1461966615416_1252" data-reactid=".5aqlq823uo.1.0.0.0.4.1.0.0.$1">Balancing the joint production of multiple ecosystem services, also referred to as the ‘multifunctionality’ of an ecosystem or landscape, requires understanding of the ecological processes that produce and economic processes that evaluate those services. Here, we review the ecological tradeoffs and compatibilities among ecosystem processes that influence ecosystem multifunctionality with respect to ecosystem services, including variation in functional strategies, constraints on community assembly and direct effects of the abiotic environment. We then review how different valuation methods may alter the magnitude of tradeoffs and compatibilities in monetary terms. Among communities, functional diversity increases ecosystem multifunctionality, but community-average trait values are emerging as important drivers of ecosystem services with greater potential to produce tradeoffs when compared to functional diversity. However, research that links organismal functional strategies to community assembly rules in real, heterogeneous landscapes demonstrate that predictable tradeoffs among species do not consistently scale up to the community level, necessitating further research on trait-based community assembly in order to develop general predictive models of biotic effects on ecosystem multifunctionality. Abiotic factors are frequently incorporated into mapping assessments of multifunctionality, but the emergent tradeoffs and compatibilities in ecosystem services driven by those factors are rarely assessed, despite a number of studies that have demonstrated their clear importance in ecosystem multifunctionality. Finally, while a variety of valuation methods are used to quantify the joint production of ecosystem services, only provisioning services are typically directly valued and assumed to have fixed correlations with other ecosystem services that can lead to inaccurate valuation, and potentially inappropriate prioritisation, of multiple ecosystem services.</span></span></span>
Cao Y, Jones C, Cuevas-Vicenttín V, Jones MB, Ludäscher B, McPhillips T, Missier P, Schwalm C, Slaughter P, Vieglais D, Walker L, Wei Y (2016) DataONE: A Data Federation with Provenance Support. Provenance and Annotation of Data and Processes: 6th International Provenance and Annotation Workshop, IPAW 2016, McLean, VA, USA, June 7-8, 2016, Proceedings 230-234.
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Read PublicationDataONE is a federated data network focusing on earth and environmental science data. We present the provenance and search features of DataONE by means of an example involving three earth scientists who interact through a DataONE Member Node. DataONE provenance systems enable reproducible research and facilitate proper attribution of scientific results transitively across generations of derived data products.
Compson ZG, Hungate BA, Whitham TG, Meneses N, Busby PE, Wojtowicz T, Ford AC, Adams KJ, Marks JC (2016) Plant genotype influences aquatic-terrestrial ecosystem linkages through timing and composition of insect emergence. Ecosphere 7 (5).
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Read PublicationTerrestrial leaf litter provides aquatic insects with an energy source and habitat structure, and species differences in litter can influence aquatic insect emergence. Emerging insects also provide energy to riparian predators. We hypothesized that plant genetics would influence the composition and timing of emerging insect communities among individual genotypes of Populus angustifolia varying in litter traits. We also compared the composition and timing of emerging insect communities on litter from mixed genotypes of three cross types of a hybridizing cottonwood complex: P. angustifolia, P. fremontii, and their F1 hybrids. Using litter harvested from an experimental common garden, we measured emerging insect community composition, abundance, and production for 12 weeks in large litter packs affixed with emergence traps. Five major findings emerged. (1) In support of the genetic similarity hypothesis, we found that, among P. angustifolia tree genotypes, litter from more closely related genotypes had more similar litter thickness, nitrogen concentrations, decomposition rates, and emerging insect communities. (2) Genetic similarity was not correlated with other litter traits, although the litter fungal community was a strong predictor of emerging insect communities. (3) Litter decomposition rate, which was the strongest predictor of emerging aquatic insect communities, was influenced by litter thickness, litter N, and the litter fungal community. (4) In contrast to strong community composition differences among P. angustifolia genotypes, differences in community composition between P. fremontii and P. angustifolia were only marginally significant, and communities on F1 hybrids were indistinguishable from P. angustifolia despitegenetic and litter trait differences. (5) Mixed litter packs muted the genetic effects observed in litter packs consisting of single genotypes. These results demonstrate that the genetic structure of riparian forests can affect the composition and timing of aquatic insect emergence. Because many riparian trees are clonal, including P. angustifolia, large clone size is likely to result in patches of genetically structured leaf litter that may influence the timing and composition of insect emergence within watersheds. Riparian restoration efforts incorporating different tree genotypes could also influence the biodiversity of emerging aquatic insects. Our work illustrates the importance of plant genes for community and ecosystem processes in riparian corridors.
Crews TE, Blesh J, Culman SW, Hayes RC, Jensen EC, Mack MC, Peoples MB, Schipanski ME (2016) Going where no grains have gone before: From early to mid-succession. Agriculture, Ecosystems & Environment 223: 223-238.
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Read PublicationAnnual-based arable agroecosystems experience among the greatest frequency, extent and magnitude of disturbance regimes of all terrestrial ecosystems. In order to control non-crop vegetation, farmers implement tillage practices and/or utilize herbicides. These practices effectively shift the farmed ecosystems to early stages of secondary succession where they remain as long as annual crops are grown. Humanity’s long-standing dependence on a disturbance-based food and fiber producing ecosystem has resulted in degraded soil structure, unsustainable levels of soil erosion, losses of soil organic matter, low nutrient and water retention, severe weed challenges, and a less-diverse or functional soil microbiome. While no-till cropping systems have reduced some hazards like soil erosion, they remain compromised with respect to ecosystem functions like water and nutrient uptake, and carbon sequestration compared to many later successional ecosystems. Recent advances in the development of perennial grain crop species invite consideration of the ecological implications of farming grains further down the successional gradient than ever before possible. In this review, we specifically explore how the nitrogen (N) economy of a mid-successional agroecosystem might differ from early-successional annual grain ecosystems as well as native mid-successional grassland ecosystems. We present a conceptual model that compares changes in soil organic matter, net ecosystem productivity, N availability, and N retention through ecosystem succession. Research from the agronomic and ecological literatures suggest that mid-successional grain agriculture should feature several ecological functions that could greatly improve synchrony between soil N supply and crop demands; these include larger active soil organic matter pools, a more trophically complex and stable soil microbiome that facilitates higher turnover rates of available N, greater N retention due to greater assimilation and seasonal translocation by deeply rooted perennial species as well as greater microbial immobilization. Compared to native mid-successional grasslands that cycle the majority of N required to maintain productivity within the ecosystem, a mid-successional agriculture would require greater external N inputs to balance N exports in food. Synthetic N fertilizer could make up this deficit, but in the interest of maximizing ecological intensification in order to minimize inputs and associated environmental consequences, we explore making up the N deficit with biological N<sub>2</sub> fixation. The dominant approach to addressing problems in agriculture is to target specific shortcomings such as nutrient retention or weed invasion. Moving agriculture down the successional gradient promises to change the nature of the ecosystem itself, shifting attention from symptom to cause, such that ecological intensification and provision of a broader suite of ecosystem services happen not in spite of, but as a consequence of agriculture.
Dong X, Grimm N, Ogle K (2016) Temporal variability in hydrology modifies the influence of geomorphology on wetland distribution along a desert stream. Journal of Ecology 104(1): 18-30.
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Read PublicationBoth geomorphic setting and dynamic environmental variables influence riverine wetland vegetation distributions. Most studies of species distributions in riverine systems emphasize either hydrological variability or geomorphic controls, but rarely consider the interaction between the two. It is unknown whether and to what extent the relationship between the geomorphic template and species distribution is modified by fluctuating environmental conditions.
This study examines how spatial patterns of riverine wetlands in a desert stream change in response to environmental shifts brought about by interannual variability in the hydrologic regime. We surveyed wetland spatial distribution and measured its abundance every June over 5 years (2009–2013) by recording patch size and presence/absence of five wetland plant species along the 12-km main stem of Sycamore Creek, Arizona, U.S.A. The study period encompassed a very large flood in January 2010, a wet year (2010), two average years (2009 and 2013) and two extremely dry years (2011 and 2012). We used a Bayesian statistical approach to analyse the relationship between geomorphic variables and wetland distribution under different hydrological conditions.
The geomorphic variables provided much greater explanatory power in dry years than in average to wet years. Hydrological conditions modified the interactions between geomorphic template and species distribution. Annual hydrological conditions affected the direction (i.e. positive or negative effect) and magnitude (i.e. the size and significance level of an effect) of these interactions, both of which gave rise to spatial patterns of wetlands. Ecosystem temporal variability, such as inter-annual and multi-year hydrological variability and longer-term ecosystem state changes, triggered complex species responses.
Synthesis. The effect of geomorphic setting on stream wetland plant distribution in this desert system is conditioned on the temporal variability in hydrology among years. Temporal transferability of the relationship between geomorphology and species distributions is therefore questionable.
Finger RA, Turetsky MR, Kielland K, Ruess RW, Mack MC, Euskirchen ES (2016) Effects of permafrost thaw on nitrogen availability and plant-soil interactions in a boreal Alaskan lowland. Journal of Ecology 104 (6): 1542-1554.
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Read PublicationIncreasing rates of permafrost thaw in boreal peatlands are converting conifer forests to waterlogged open wetlands. Permafrost thaw increases soil nitrogen (N) availability, but it is unclear whether such changes are due solely to changes in surface soil N mineralization or N mobilization from thawing permafrost soils at depth.
We examined plant species composition and N availability along triplicate permafrost thaw gradients in Alaskan peatlands. Each gradient comprised four community types including: 1) a permafrost peatland with intact permafrost; 2) a drunken forest experiencing active thaw; 3) a moat representing initial complete thaw; and 4) a collapse scar bog representing several decades of post-thaw succession.
Concentrations of dissolved organic (DON) and inorganic N (DIN) in the upper 60 cm of soil increased along the permafrost thaw gradients. The drunken forest had the greatest mean concentrations of total dissolved N relative to the other community types, primarily due to greater concentrations of large molecular DON. The moat and collapse bog had significantly greater inorganic N concentrations than the permafrost or drunken forest, suggesting that changes in N availability are not a short-term effect, but can be sustained for decades or centuries. Across all plant community types, DIN and DON concentrations increased with soil depth during maximum seasonal ice thaw (September), suggesting that deeper soil horizons are important reservoirs of N post-thaw.
Vegetation responses to permafrost thaw included changes in plant community composition shifting from upland forest species to hydrophilic vegetation with deeper rooting profiles in the collapse scar bogs, and changes in foliar N and δ15N values. N concentrations in plant foliage and litterfall increased with concentrations of DIN during collapse bog succession, suggesting that plants are utilizing additional mineralized N.
Synthesis: Our results suggest that the conversion of forest to wetlands associated with permafrost thaw in boreal lowlands increases N availability, at least in part by increasing turnover of deep soil organic matter. Plants appear to utilize these additional deeper N sources over timescales of years to centuries following permafrost thaw.
Fisher JB, Sikka M, Huntzinger DN, Schwalm C, Liu J (2016) Technical note: 3-hourly temporal downscaling of monthly global terrestrial biosphere model net ecosystem exchange. Biogeosciences 13(14): 4271-4277.
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Read PublicationThe land surface provides a boundary condition to atmospheric forward and flux inversion models. These models require prior estimates of CO<sub>2</sub> fluxes at relatively high temporal resolutions (e.g., 3-hourly) because of the high frequency of atmospheric mixing and wind heterogeneity. However, land surface model CO<sub>2</sub> fluxes are often provided at monthly time steps, typically because the land surface modeling community focuses more on time steps associated with plant phenology (e.g., seasonal) than on sub-daily phenomena. Here, we describe a new dataset created from 15 global land surface models and 4 ensemble products in the Multi-scale Synthesis and Terrestrial Model Intercomparison Project (MsTMIP), temporally downscaled from monthly to 3-hourly output. We provide 3-hourly output for each individual model over 7 years (2004–2010), as well as an ensemble mean, a weighted ensemble mean, and the multi-model standard deviation. Output is provided in three different spatial resolutions for user preferences: 0.5° × 0.5°, 2.0° × 2.5°, and 4.0° × 5.0° (latitude × longitude).
Gehring CA, Hayer M, Flores-Rentería L, Krohn AF, Schwartz E, Dijkstra P (2016) Cheatgrass invasion alters the abundance and composition of dark septate fungal communities in sagebrush steppe. Botany 94 (6): 481-491.
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Read PublicationInvasive, non-native plant species can alter soil microbial communities in ways that contribute to their persistence. While most studies emphasize mycorrhizal fungi, invasive plants also may influence communities of dark septate fungi (DSF), common root endophytes that can function like mycorrhizas. We tested the hypothesis that a widespread invasive plant in the western United States, cheatgrass (Bromus tectorum), influenced the abundance and community composition of DSF by examining the roots and rhizosphere soils of cheatgrass and two native plant species in cheatgrass invaded and non-invaded areas of sagebrush steppe. We focused on cheatgrass because it is negatively affected by mycorrhizal fungi and colonized by DSF. We found that DSF root colonization and operational taxonomic (OTU) richness were significantly higher in sagebrush (Artemisia tridentata) and rice grass (Achnatherum hymenoides) from invaded areas than non-invaded areas. Cheatgrass roots had similar levels of DSF colonization and OTU richness as native plants. The community composition of DSF varied with invasion in the roots and soils of native species and among the roots of the three plant species in invaded areas. The substantial changes in DSF we observed following cheatgrass invasion argue for comparative studies of DSF function in native and non-native plant species.
Grosse G, Goetz S, McGuire AD, Romanovsky VE and Schuur EAG (2016) Changing permafrost in a warming world and feedbacks to the Earth system. Environmental Research Letters 11(4): 040201.
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Read Publication<span id="yui_3_14_1_1_1461105258100_1608" class="foldable-text" data-reactid=".qoiwja49og.1.0.0.0.4.1.0"><span id="yui_3_14_1_1_1461105258100_1773" class="text-with-line-breaks" data-reactid=".qoiwja49og.1.0.0.0.4.1.0.0"><span id="yui_3_14_1_1_1461105258100_1772" data-reactid=".qoiwja49og.1.0.0.0.4.1.0.0.$1">The permafrost component of the cryosphere is changing dramatically, but the permafrost region is not well monitored and the consequences of change are not well understood. Changing permafrost interacts with ecosystems and climate on various spatial and temporal scales. The feedbacks resulting from these interactions range from local impacts on topography, hydrology, and biology to complex influences on global scale biogeochemical cycling. This review contributes to this focus issue by synthesizing its 28 multidisciplinary studies which provide field evidence, remote sensing observations, and modeling results on various scales. We synthesize study results from a diverse range of permafrost landscapes and ecosystems by reporting key observations and modeling outcomes for permafrost thaw dynamics, identifying feedbacks between permafrost and ecosystem processes, and highlighting biogeochemical feedbacks from permafrost thaw. We complete our synthesis by discussing the progress made, stressing remaining challenges and knowledge gaps, and providing an outlook on future needs and research opportunities in the study of permafrost–ecosystem–climate interactions.</span></span></span>
Harrison JG, Forister ML, Parchman TL, Koch GW (2016) Vertical stratification of the foliar fungal community in the world’s tallest trees. American Journal of Botany 103 (12): 2087 - 2095.
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<p id="p-1"><strong>PREMISE OF THE STUDY:</strong> The aboveground tissues of plants host numerous, ecologically important fungi, yet patterns in the spatial distribution of these fungi remain little known. Forest canopies in particular are vast reservoirs of fungal diversity, but intracrown variation in fungal communities has rarely been explored. Knowledge of how fungi are distributed throughout tree crowns will contribute to our understanding of interactions between fungi and their host trees and is a first step toward investigating drivers of community assembly for plant-associated fungi. Here we describe spatial patterns in fungal diversity within crowns of the world’s tallest trees, coast redwoods (<em>Sequoia sempervirens</em>).</p>
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<p id="p-2"><strong>METHODS:</strong> We took a culture-independent approach, using the Illumina MiSeq platform, to characterize the fungal assemblage at multiple heights within the crown across the geographical range of the coast redwood.</p>
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<p id="p-3"><strong>KEY RESULTS:</strong> Within each tree surveyed, we uncovered evidence for vertical stratification in the fungal community; different portions of the tree crown harbored different assemblages of fungi. We also report between-tree variation in the fungal community within redwoods.</p>
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<p id="p-4"><strong>CONCLUSIONS:</strong> Our results suggest the potential for vertical stratification of fungal communities in the crowns of other tall tree species and should prompt future study of the factors giving rise to this stratification.</p>
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Hayer M, Schwartz E, Marks JC, Koch BJ, Morrissey EM, Schuettenberg AA, Hungate BA (2016) Identification of growing bacteria during litter decomposition in freshwater through H2 18O quantitative stable isotope probing. Environmental Microbiology Reports 8(6): 975-982.
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Read PublicationIdentification of microorganisms that facilitate the cycling of nutrients in freshwater is paramount to understanding how these ecosystems function. Here, we identify growing aquatic bacteria using <span class="math-equation-construct" data-equation-construct="true"><span class="math-equation-image" data-equation-image="true"><img class="inlineGraphic" src="http://onlinelibrary.wiley.com/store/10.1111/1758-2229.12475/asset/equation/emi412475-math-0002.png?v=1&s=800650689e2f28b7ca342149c1b25838931bf5b9" alt="math formula" /></span></span> quantitative stable isotope probing. During 8 day incubations in 97 atom %<span class="math-equation-construct" data-equation-construct="true"><span class="math-equation-image" data-equation-image="true"><img class="inlineGraphic" src="http://onlinelibrary.wiley.com/store/10.1111/1758-2229.12475/asset/equation/emi412475-math-0003.png?v=1&s=b511335828cf868c20f2f0a4a86a7b4b83a09dc9" alt="math formula" /></span></span>, 54% of the taxa grew. The most abundant phyla among growing taxa were <em>Proteobacteria</em> (45%), <em>Bacteroidetes</em> (30%) and <em>Firmicutes</em> (10%). Taxa differed in isotopic enrichment, reflecting variation in DNA replication of bacterial populations. At the class level, the highest atom fraction excess was observed for OPB41 and δ-<em>Proteobacteria</em>. There was no linear relationship between <sup>18</sup>O incorporation and abundance of taxa. δ-<em>Proteobacteria</em> and OPB41 were not abundant, yet the DNA of both taxa was highly enriched in <sup>18</sup>O. <em>Bacteriodetes</em>, in contrast, were abundant but not highly enriched. Our study shows that a large proportion of the bacterial taxa found on decomposing leaf litter grew slowly, and several low abundance taxa were highly enriched. These findings indicating that rare organisms may be important for the decomposition of leaf litter in streams, and that quantitative stable isotope probing with <span class="math-equation-construct" data-equation-construct="true"><span class="math-equation-image" data-equation-image="true"><img class="inlineGraphic" src="http://onlinelibrary.wiley.com/store/10.1111/1758-2229.12475/asset/equation/emi412475-math-0004.png?v=1&s=1fde9faebe8b6460b636559f2fe2464644157799" alt="math formula" /></span></span> can be used to advance our understanding of microorganisms in freshwater by identifying species that are growing in complex communities.
Hewitt RE , Hollingsworth TN, Chapin III FS, and Taylor DL (2016) Fire‑severity effects on plant–fungal interactions after a novel tundra wildfire disturbance: implications for arctic shrub and tree migration. BMC Ecology 16(1): 25.
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Read Publication<div id="yui_3_14_1_1_1463605085348_1720" class="selectable" data-canvas-width="91.9909420168124"><strong>Background:</strong> Vegetation change in high latitude tundra ecosystems is expected to accelerate due to increased wild-fire activity. High-severity fires increase the availability of mineral soil seedbeds, which facilitates recruitment, yet fire also alters soil microbial composition, which could significantly impact seedling establishment.</div>
<div id="yui_3_14_1_1_1463605085348_1733" class="selectable" data-canvas-width="56.02443236414706"><strong>Results:</strong> We investigated the effects of fire severity on soil biota and associated effects on plant performance for two plant species predicted to expand into Arctic tundra. We inoculated seedlings in a growth chamber experiment with soils collected from the largest tundra fire recorded in the Arctic and used molecular tools to characterize root-associated fungal communities. Seedling biomass was significantly related to the composition of fungal inoculum. Biomass decreased as fire severity increased and the proportion of pathogenic fungi increased.</div>
<div class="selectable" data-canvas-width="91.47017517924458"><strong>Conclusions:</strong> Our results suggest that effects of fire severity on soil biota reduces seedling performance and thus we hypothesize that in certain ecological contexts fire-severity effects on plant–fungal interactions may dampen the expected increases in tree and shrub establishment after tundra fire.</div>
<div class="selectable" data-canvas-width="75.30960428439917"><strong>Keywords:</strong> Alnus viridis, Arctic tundra, ARISA, Climate change, Fire severity, Fungal internal transcribed spacer (ITS), Picea mariana, Shrub expansion, Treeline</div>
Hewitt RE, Bennett AP, Breen AL, Hollingsworth TN, Taylor DL, Chapin FS, Rupp TS (2016) Getting to the root of the matter: Landscape implications of plant-fungal interactions for tree migration in Alaska. Landscape Ecology 31(4): 895-911.
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<h3 class="Heading">Context</h3>
<p id="Par1" class="Para">Forecasting the expansion of forest into Alaska tundra is critical to predicting regional ecosystem services, including climate feedbacks such as carbon storage. Controls over seedling establishment govern forest development and migration potential. Ectomycorrhizal fungi (EMF), obligate symbionts of all Alaskan tree species, are particularly important to seedling establishment, yet their significance to landscape vegetation change is largely unknown.</p>
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<h3 class="Heading">Objective</h3>
<p id="Par2" class="Para">We used ALFRESCO, a landscape model of wildfire and vegetation dynamics, to explore whether EMF inoculum potential influences patterns of tundra afforestation and associated flammability.</p>
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<h3 class="Heading">Methods</h3>
<p id="Par3" class="Para">Using two downscaled CMIP3 general circulation models (ECHAM5 and CCCMA) and a mid-range emissions scenario (A1B) at a 1 km<sup>2</sup> resolution, we compared simulated tundra afforestation rates and flammability from four parameterizations of EMF effects on seedling establishment and growth from 2000 to 2100.</p>
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<h3 class="Heading">Results</h3>
<p id="Par4" class="Para">Modeling predicted an 8.8–18.2 % increase in forest cover from 2000 to 2100. Simulations that explicitly represented landscape variability in EMF inoculum potential showed a reduced percent change afforestation of up to a 2.8 % due to low inoculum potential limiting seedling growth. This reduction limited fuel availability and thus, cumulative area burned. Regardless of inclusion of EMF effects in simulations, landscape flammability was lower for simulations driven by the wetter and cooler CCCMA model than the warmer and drier ECHAM5 model, while tundra afforestation was greater.</p>
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<h3 class="Heading">Conclusions</h3>
<p id="Par5" class="Para">Results suggest abiotic factors are the primary driver of tree migration. Simulations including EMF effects, a biotic factor, yielded more conservative estimates of land cover change across Alaska that better-matched empirical estimates from the previous century.</p>
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Hinckley E-L, Bonan G, Bowen G, Colman B, Duffy P, Goodale C, Houlton B, Marín-Spiotta E, Ogle K, Ollinger S, Paul E, Vitousek P, Weathers K, Williams D (2016) The soil and plant biogeochemistry sampling design for The National Ecological Observatory Network. Ecosphere 7(3): e01234.
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Read PublicationHuman impacts on biogeochemical cycles are evident around the world, from changes to forest structure and function due to atmospheric deposition, to eutrophication of surface waters from agricultural effluent, and increasing concentrations of carbon dioxide (CO<sub>2</sub>) in the atmosphere. The National Ecological Observatory Network (NEON) will contribute to understanding human effects on biogeochemical cycles from local to continental scales. The broad NEON biogeochemistry measurement design focuses on measuring atmospheric deposition of reactive mineral compounds and CO<sub>2</sub> fluxes, ecosystem carbon (C) and nutrient stocks, and surface water chemistry across 20 eco-climatic domains within the United States for 30 yr. Herein, we present the rationale and plan for the ground-based measurements of C and nutrients in soils and plants based on overarching or “high-level” requirements agreed upon by the National Science Foundation and NEON. The resulting design incorporates early recommendations by expert review teams, as well as recent input from the larger natural sciences community that went into the formation and interpretation of the requirements, respectively. NEON's efforts will focus on a suite of data streams that will enable end-users to study and predict changes to biogeochemical cycling and transfers within and across air, land, and water systems at regional to continental scales. At each NEON site, there will be an initial, one-time effort to survey soil properties to 1 m (including soil texture, bulk density, pH, baseline chemistry) and vegetation community structure and diversity. A sampling program will follow, focused on capturing long-term trends in soil C, nitrogen (N), and sulfur stocks, isotopic composition (of C and N), soil N transformation rates, phosphorus pools, and plant tissue chemistry and isotopic composition (of C and N). To this end, NEON will conduct extensive measurements of soils and plants within stratified random plots distributed across each site. The resulting data will be a new resource for members of the scientific community interested in addressing questions about long-term changes in continental-scale biogeochemical cycles, and is predicted to inspire further process-based research.
Hungate BA, Kearns DN, Ogle K, Caron M, Marks JC, Rogg HW (2016) Hydrogen Isotopes as a Sentinel of Biological Invasion by the Japanese Beetle, Popillia japonica (Newman). Plos One 11(3): e0149599.
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Read PublicationInvasive species alter ecosystems, threaten native and endangered species, and have negative economic impacts. Knowing where invading individuals are from and when they arrive to a new site can guide management. Here, we evaluated how well the stable hydrogen isotope composition (δ2H) records the recent origin and time since arrival of specimens of the invasive Japanese beetle (Popillia japonica Newman) captured near the Portland International Airport (Oregon, U.S.A.). The δ2H of Japanese beetle specimens collected from sites across the contiguous U.S.A. reflected the δ2H of local precipitation, a relationship similar to that documented for other organisms, and one confirming the utility of δ2H as a geographic fingerprint. Within weeks after experimental relocation to a new isotopic environment, the δ2H of beetles changed linearly with time, demonstrating the potential for δ2H to also mark the timing of arrival to a new location. We used a hierarchical Bayesian model to estimate the recent geographical origin and timing of arrival of each specimen based on its δ2H value. The geographic resolution was broad, with values consistent with multiple regions of origin in the eastern U.S.A., slightly favoring the southeastern U.S.A. as the more likely source. Beetles trapped from 2007–2010 had arrived 30 or more days prior to trapping, whereas the median time since arrival declined to 3–7 days for beetles trapped from 2012–2014. This reduction in the time between arrival and trapping at the Portland International Airport supports the efficacy of trapping and spraying to prevent establishment. More generally, our analysis shows how stable isotopes can serve as sentinels of biological invasions, verifying the efficacy of control measures, or, alternatively, indicating when those measures show signs of failure.
Hurteau MD, Liang S, Martin KL, North MP, Koch GW, and Hungate BA (2016) Restoring forest structure and process stabilizes forest carbon in wildfire-prone southwestern ponderosa pine forests. Ecological Applications 26(2): 382–391.
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Read PublicationChanging climate and a legacy of fire-exclusion have increased the probability of high-severity wildfire, leading to an increased risk of forest carbon loss in ponderosa pine forests in the southwestern USA. Efforts to reduce high-severity fire risk through forest thinning and prescribed burning require both the removal and emission of carbon from these forests, and any potential carbon benefits from treatment may depend on the occurrence of wildfire. We sought to determine how forest treatments alter the effects of stochastic wildfire events on the forest carbon balance. We modeled three treatments (control, thin-only, and thin and burn) with and without the occurrence of wildfire. We evaluated how two different probabilities of wildfire occurrence, 1% and 2% per year, might alter the carbon balance of treatments. In the absence of wildfire, we found that thinning and burning treatments initially reduced total ecosystem carbon (TEC) and increased net ecosystem carbon balance (NECB). In the presence of wildfire, the thin and burn treatment TEC surpassed that of the control in year 40 at 2%/yr wildfire probability, and in year 51 at 1%/yr wildfire probability. NECB in the presence of wildfire showed a similar response to the no-wildfire scenarios: both thin-only and thin and burn treatments increased the C sink. Treatments increased TEC by reducing both mean wildfire severity and its variability. While the carbon balance of treatments may differ in more productive forest types, the carbon balance benefits from restoring forest structure and fire in southwestern ponderosa pine forests are clear.
Ito A, Inatomi M, Huntzinger DN, Schwalm CR, Michalak AM, Cook R, King AW, Mao J, Wei Y, Mac Post W, Wang W (2016) DECADAL TRENDS IN THE SEASONAL-CYCLE AMPLITUDE OF TERRESTRIAL CO2 EXCHANGE RESULTING FROM THE ENSEMBLE OF TERRESTRIAL BIOSPHERE MODELS. Tellus 68: 28968.
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Read PublicationThe seasonal-cycle amplitude (SCA) of the atmosphere–ecosystem carbon dioxide (CO<sub>2</sub>) exchange rate is a useful metric of the responsiveness of the terrestrial biosphere to environmental variations. It is unclear, however, what underlying mechanisms are responsible for the observed increasing trend of SCA in atmospheric CO<sub>2</sub> concentration. Using output data from the Multi-scale Terrestrial Model Intercomparison Project (MsTMIP), we investigated how well the SCA of atmosphere–ecosystem CO<sub>2</sub> exchange was simulated with 15 contemporary terrestrial ecosystem models during the period 1901–2010. Also, we made attempt to evaluate the contributions of potential mechanisms such as atmospheric CO<sub>2</sub>, climate, land-use, and nitrogen deposition, through factorial experiments using different combinations of forcing data. Under contemporary conditions, the simulated global-scale SCA of the cumulative net ecosystem carbon flux of most models was comparable in magnitude with the SCA of atmospheric CO<sub>2</sub> concentrations. Results from factorial simulation experiments showed that elevated atmospheric CO<sub>2</sub> exerted a strong influence on the seasonality amplification. When the model considered not only climate change but also land-use and atmospheric CO<sub>2</sub>changes, the majority of the models showed amplification trends of the SCAs of photosynthesis, respiration, and net ecosystem production (+0.19 % to +0.50 % yr<sup>−1</sup>). In the case of land-use change, it was difficult to separate the contribution of agricultural management to SCA because of inadequacies in both the data and models. The simulated amplification of SCA was approximately consistent with the observational evidence of the SCA in atmospheric CO<sub>2</sub> concentrations. Large inter-model differences remained, however, in the simulated global tendencies and spatial patterns of CO<sub>2</sub> exchanges. Further studies are required to identify a consistent explanation for the simulated and observed amplification trends, including their underlying mechanisms. Nevertheless, this study implied that monitoring of ecosystem seasonality would provide useful insights concerning ecosystem dynamics.
Johnston ER, Rodríguez-R LM, Luo C, Yuan MM, Wu L, He Z, Schuur EA, Luo Y, Tiedje JM, Zhou J, Konstantinidis KT (2016) Metagenomics reveals pervasive bacterial populations and reduced community diversity across the Alaska tundra ecosystem. Frontiers in Microbiology 7: 579.
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Read Publication<div id="yui_3_14_1_1_1461088040208_3844" class="selectable" data-canvas-width="886.7532679738565">How soil microbial communities contrast with respect to taxonomic and functional composition within and between ecosystems remains an unresolved question that is central to predicting how global anthropogenic change will affect soil functioning and services. In particular, it remains unclear how small-scale observations of soil communities based on the typical volume sampled (1-2 grams) are generalizable to ecosystem- scale responses and processes. This is especially relevant for remote, northern latitude soils, which are challenging to sample and are also thought to be more vulnerable to climate change compared to temperate soils. Here, we employed well replicated shotgun metagenome and 16S rRNA gene amplicon sequencing to characterize community composition and metabolic potential in Alaskan tundra soils, combining our own datasets with those publically available from distant tundra and temperate grassland and agriculture habitats. We found that the abundance of many taxa and metabolic functions differed substantially between tundra soil metagenomes relative to those from temperate soils, and that a high degree of OTU-sharing exists between tundra locations. Tundra soils were an order of magnitude less complex than their temperate counterparts, allowing for near- complete coverage of microbial community richness (~92% breadth) by sequencing, and the recovery of twenty-seven high-quality, almost complete (>80% completeness) population bins. These population bins, collectively, made up to ~10% of the metagenomic datasets, and represented diverse taxonomic groups and metabolic lifestyles tuned toward sulfur cycling, hydrogen metabolism, methanotrophy, and organic matter oxidation. Several population bins, including members of Acidobacteria, Actinobacteria, and Proteobacteria, were also present in geographically distant (~100-530 km apart) tundra habitats (full genome representation and up to 99.6% genome-derived average nucleotide identity). Collectively, our results revealed that Alaska tundra microbial communities are less diverse and more homogenous across spa tial scales than previously anticipated, and provided DNA sequencesof abundant populations and genes that would be relevant for future studies of the effects of environmental change on tundra ecosystems.</div>
Kennedy TA, Muehlbauer JD, Yackulic CB,Lytle DA, Miller SW, Dibble KW, Kortenhoeven EW, Metcalfe AN, Baxter CV (2016) Flow Management for Hydropower Extirpates Aquatic Insects, Undermining River Food Webs. BioScience 66(7): 561-575.
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Read PublicationDams impound the majority of rivers and provide important societal benefits, especially daily water releases that enable on-peak hydroelectricity generation. Such “hydropeaking” is common worldwide, but its downstream impacts remain unclear. We evaluated the response of aquatic insects, a cornerstone of river food webs, to hydropeaking using a life history–hydrodynamic model. Our model predicts that aquatic-insect abundance will depend on a basic life-history trait—adult egg-laying behavior—such that open-water layers will be unaffected by hydropeaking, whereas ecologically important and widespread river-edge layers, such as mayflies, will be extirpated. These predictions are supported by a more-than-2500-sample, citizen-science data set of aquatic insects from the Colorado River in the Grand Canyon and by a survey of insect diversity and hydropeaking intensity across dammed rivers of the Western United States. Our study reveals a hydropeaking-related life history bottleneck that precludes viable populations of many aquatic insects from inhabiting regulated rivers.
Kranabetter MJ, McLaughlan KK, Enders SK, Fraterrigo JM, Higuera PE, Morris JL, Rastetter EN, Barnes R, Buma B, Gavin DG, Gerhart LM, Gilson L, Hietz P, Mack MC, McNeil B, Perakis S (2016) A Framework to Assess Biogeochemical Response to Ecosystem Disturbance Using Nutrient Partitioning Ratios. Ecosystems 19(3): 387–395.
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Read PublicationDisturbances affect almost all terrestrial ecosystems, but it has been difficult to identify general principles regarding these influences. To improve our understanding of the long-term consequences of disturbance on terrestrial ecosystems, we present a conceptual framework that analyzes disturbances by their biogeochemical impacts. We posit that the ratio of soil and plant nutrient stocks in mature ecosystems represents a characteristic site property. Focusing on nitrogen (N), we hypothesize that this partitioning ratio (soil N: plant N) will undergo a predictable trajectory after disturbance. We investigate the nature of this partitioning ratio with three approaches: (1) nutrient stock data from forested ecosystems in North America, (2) a process-based ecosystem model, and (3) conceptual shifts in site nutrient availability with altered disturbance frequency. Partitioning ratios could be applied to a variety of ecosystems and successional states, allowing for improved temporal scaling of disturbance events. The generally short-term empirical evidence for recovery trajectories of nutrient stocks and partitioning ratios suggests two areas for future research. First, we need to recognize and quantify how disturbance effects can be accreting or depleting, depending on whether their net effect is to increase or decrease ecosystem nutrient stocks. Second, we need to test how altered disturbance frequencies from the present state may be constructive or destructive in their effects on biogeochemical cycling and nutrient availability. Long-term studies, with repeated sampling of soils and vegetation, will be essential in further developing this framework of biogeochemical response to disturbance.
Kropp H, Ogle K, Wojciechowski MF (2016) A framework for partitioning plant rooting profiles from neighbours using multiple data types. Journal of Vegetation Science 27(3): 587-595.
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Read Publication<section id="jvs12377-sec-0001" class="article-section article-body-section">
<h3>Aim</h3>
Vertical root distributions (‘profiles’) influence plant water use and productivity, and the differentiation of root profiles between neighbouring species can indicate the degree of plant interactions and niche partitioning. However, quantifying multiple species' root distributions in the field can be labour intensive and highly destructive to the soil and plants. We describe a method for partitioning multiple species roots using minimally destructive methods to determine if neighbour interactions alter the root profile of a common desert shrub, <em>Larrea tridentata</em>(creosote bush).
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<h3>Location</h3>
Sonoran Desert, central Arizona, USA.
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<h3>Methods</h3>
We obtained root and soil samples from soil cores collected around <em>Larrea</em> growing alone and next to three different neighbouring species. Bulk root mass was measured for each soil sample, and <em>Larrea</em> and neighbouring species root presence was determined with molecular identification methods. Water extracted from the soil and paired stem samples was analysed for its stable isotope composition (D and <sup>18</sup>O). Species-specific (i.e. <em>Larrea</em> and neighbouring species) root biomass and fractional active root area were estimated through a hierarchical statistical modelling approach that combined all three data sets and accounted for detection errors.
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<h3>Results</h3>
The combined data model successfully partitioned <em>Larrea</em> root biomass from neighbouring plants and provided biologically relevant estimates of rooting profiles with greater certainty than individual analyses of each data source. The data model results indicate that plant neighbours alter <em>Larrea</em>'s root profile; <em>Larrea</em> growing under tree species had significantly higher root biomass in shallow soil layers than <em>Larre</em>a growing alone.
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<h3>Conclusions</h3>
Our framework requires minimally destructive sampling methods, and accounts for sampling errors associated with different methods. We demonstrate the utility of our approach with a common desert shrub species, which illustrated that plant neighbours can alter the <em>Larrea</em>vertical root profile. Our approach is useful in problematic study systems fraught with sample collection issues or supporting species with inhibitory compounds that prohibit the use of more sophisticated molecular methods to identify the presence of other species' roots.
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Kwon MJ, Heimann M, Kolle O, Luus KA, Schuur EAG, Zimov N, Zimov SA, Göckede M. (2016) Long-term drainage reduces CO2 uptake and increases CO2 emission on a Siberian floodplain due to shifts in vegetation community and soil thermal characteristics. Biogeosciences 13(14): 4219-4235.
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Read Publication<span id="yui_3_14_1_1_1470075122251_2130" class="foldable-text" data-reactid="113"><span class="text-with-line-breaks">With increasing air temperatures and changing precipitation patterns forecast for the Arctic over the coming decades, the thawing of ice-rich permafrost is expected to increasingly alter hydrological conditions by creating mosaics of wetter and drier areas. The objective of this study is to investigate how 10 years of lowered water table depths of wet floodplain ecosystems would affect CO2 fluxes measured using a closed chamber system, focusing on the role of long-term changes in soil thermal characteristics and vegetation community structure. Drainage diminishes the heat capacity and thermal conductivity of organic soil, leading to warmer soil temperatures in shallow layers during the daytime and colder soil temperatures in deeper layers, resulting in a reduction in thaw depths. These soil temperature changes can intensify growing-season heterotrophic respiration by up to 95 %. With decreased autotrophic respiration due to reduced gross primary production under these dry conditions, the differences in ecosystem respiration rates in the present study were 25%. We also found that a decade-long drainage installation significantly increased shrub abundance, while decreasing Eriophorum angustifolium abundance resulted in Carex sp. dominance. These two changes had opposing influences on gross primary production during the growing season: while the increased abundance of shrubs slightly increased gross primary production, the replacement of E. angustifolium by Carex sp. significantly decreased it. With the effects of ecosystem respiration and gross primary production combined, net CO2 uptake rates varied between the two years, which can be attributed to Carex-dominated plots' sensitivity to climate. However, underlying processes showed consistent patterns: 10 years of drainage increased soil temperatures in shallow layers and replaced E. angustifolium by Carex sp., which increased CO2 emission and reduced CO2 uptake rates. During the non-growing season, drainage resulted in 4 times more CO2 emissions, with high sporadic fluxes; these fluxes were induced by soil temperatures, E. angustifolium abundance, and air pressure.</span></span>
Laflower DM, Hurteau MD, Koch GW, North MP, Hungate BA (2016) Climate-driven changes in forest succession and the influence of management on forest carbon dynamics in the Puget Lowlands of Washington State, USA. Forest Ecology and Management 362: 194-204.
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Read PublicationProjecting the response of forests to changing climate requires understanding how biotic and abiotic controls on tree growth will change over time. As temperature and interannual precipitation variability increase, the overall forest response is likely to be influenced by species-specific responses to changing climate. Management actions that alter composition and density may help buffer forests against the effects of changing climate, but may require tradeoffs in ecosystem services. We sought to quantify how projected changes in climate and different management regimes would alter the composition and productivity of Puget Lowland forests in Washington State, USA. We modeled forest responses to four treatments (control, burn-only, thin-only, thin-and-burn) under five different climate scenarios: baseline climate (historical) and projections from two climate models (CCSM4 and CNRM-CM5), driven by moderate (RCP 4.5) and high (RCP 8.5) emission scenarios. We also simulated the effects of intensive management to restore Oregon white oak woodlands (Quercus garryana) for the western gray squirrel (Sciurus griseus) and quantified the effects of these treatments on the probability of oak occurrence and carbon sequestration. At the landscape scale we found little difference in carbon dynamics between baseline and moderate emission scenarios. However, by late-century under the high emission scenario, climate change reduced forest productivity and decreased species richness across a large proportion of the study area. Regardless of the climate scenario, we found that thinning and burning treatments increased the carbon sequestration rate because of decreased resource competition. However, increased productivity with management was not sufficient to prevent an overall decline in productivity under the high emission scenario. We also found that intensive oak restoration treatments were effective at increasing the probability of oak presence and that the limited extent of the treatments resulted in small declines in total ecosystem carbon across the landscape as compared to the thin-and-burn treatment. Our research suggests that carbon dynamics in this system under the moderate emission scenario may be fairly consistent with the carbon dynamics under historical climate, but that the high emission scenario may alter the successional trajectory of these forests.
Le Roux X, Bouskill NJ, Niboyet A, Barthes L, Dijkstra P, Field CB, Hungate BA, Lerondelle C, Pommier T, Tang J, Terada A, Tourna M, and Poly F (2016) Predicting the Responses of Soil Nitrite-Oxidizers to Multi-Factorial Global Change: A Trait-Based Approach. Frontiers in Microbiology 7: 628.
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Read PublicationSoil microbial diversity is huge and a few grams of soil contain more bacterial taxa than there are bird species on Earth. This high diversity often makes predicting the responses of soil bacteria to environmental change intractable and restricts our capacity to predict the responses of soil functions to global change. Here, using a long-term field experiment in a California grassland, we studied the main and interactive effects of three global change factors (increased atmospheric CO2 concentration, precipitation and nitrogen addition, and all their factorial combinations, based on global change scenarios for central California) on the potential activity, abundance and dominant taxa of soil nitrite-oxidizing bacteria (NOB). Using a trait-based model, we then tested whether categorizing NOB into a few functional groups unified by physiological traits enables understanding and predicting how soil NOB respond to global environmental change. Contrasted responses to global change treatments were observed between three main NOB functional types. In particular, putatively mixotrophic <i>Nitrobacter</i>, rare under most treatments, became dominant under the ‘High CO2CNitrogenCPrecipitation’ treatment. The mechanistic trait-based model, which simulated ecological niches of NOB types consistent with previous ecophysiological reports, helped predicting the observed effects of global change on NOB and elucidating the underlying biotic and abiotic controls. Our results are a starting point for representing the overwhelming diversity of soil bacteria by a few functional types that can be incorporated into models of terrestrial ecosystems and biogeochemical processes.
Luo Y, Ahlström A, Allison SD, Batjes NH, Brovkin V, Carvalhais N, Chappell A, Ciais P, Davidson EA, Finzi A, Georgiou K, Guenet B, Hararuk O, Harden JW, He Y, Hopkins F, Jiang L, Koven C, Jackson RB, Jones CD, Lara MJ, Liang J, McGuire AD, Parton W, Peng C, Randerson JT, Salazar A, Sierra CA, Smith MJ, Tian H, Todd-Brown KEO, Torn M, van Groenigen KJ, Wang YP, West TO, Wei Y, Wieder WR, Xia J, Xu X, Xu X, Zhou T (2016) Toward more realistic projections of soil carbon dynamics by Earth system models. Global Biogeochemical Cycles 30(1): 40-56.
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Read Publication<section id="abstract" class="article-section article-section--abstract">
<div class="article-section__content mainAbstract">
Soil carbon (C) is a critical component of Earth system models (ESMs), and its diverse representations are a major source of the large spread across models in the terrestrial C sink from the third to fifth assessment reports of the Intergovernmental Panel on Climate Change (IPCC). Improving soil C projections is of a high priority for Earth system modeling in the future IPCC and other assessments. To achieve this goal, we suggest that (1) model structures should reflect real-world processes, (2) parameters should be calibrated to match model outputs with observations, and (3) external forcing variables should accurately prescribe the environmental conditions that soils experience. First, most soil C cycle models simulate C input from litter production and C release through decomposition. The latter process has traditionally been represented by first-order decay functions, regulated primarily by temperature, moisture, litter quality, and soil texture. While this formulation well captures macroscopic soil organic C (SOC) dynamics, better understanding is needed of their underlying mechanisms as related to microbial processes, depth-dependent environmental controls, and other processes that strongly affect soil C dynamics. Second, incomplete use of observations in model parameterization is a major cause of bias in soil C projections from ESMs. Optimal parameter calibration with both pool- and flux-based data sets through data assimilation is among the highest priorities for near-term research to reduce biases among ESMs. Third, external variables are represented inconsistently among ESMs, leading to differences in modeled soil C dynamics. We recommend the implementation of traceability analyses to identify how external variables and model parameterizations influence SOC dynamics in different ESMs. Overall, projections of the terrestrial C sink can be substantially improved when reliable data sets are available to select the most representative model structure, constrain parameters, and prescribe forcing fields.
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McGuire AD, Koven C, Lawrence DM, Clein JS, Xia J, Beer C, Burke E, Chen G, Chen X, Delire C, Jafarov E, MacDougall AH, Marchenko S, Nicolsky D, Peng S, Rinke A, Saito K, Zhang W, Alkama R, Bohn TJ, Ciais P, Decharme B, Ekici A, Gouttevin I, Hajima T, Hayes DJ, Ji D, Krinner G, Lettenmaier DP, Luo Y, Miller PA, Moore JC. Romanovsky V, Schädel C, Schaefer K, Schuur EAG, Smith B, Sueyoshi T, Zhuang Q (2016) Variability in the sensitivity among model simulations of permafrost and carbon dynamics in the permafrost region between 1960 and 2009. Global Biogeochemical Cycles 30 (7): 1015-1037.
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Read PublicationA significant portion of the large amount of carbon (C) currently stored in soils of the permafrost region in the Northern Hemisphere has the potential to be emitted as the greenhouse gases CO2 and CH4 under a warmer climate. In this study we evaluated the variability in the sensitivity of permafrost and C in recent decades among land surface model simulations over the permafrost region between 1960 and 2009. The 15 model simulations all predict a loss of near-surface permafrost (within 3 m) area over the region, but there are large differences in the magnitude of the simulated rates of loss among the models (0.2 to 58.8 × 103 km2 yr−1). Sensitivity simulations indicated that changes in air temperature largely explained changes in permafrost area, although interactions among changes in other environmental variables also played a role. All of the models indicate that both vegetation and soil C storage together have increased by 156 to 954 Tg C yr−1 between 1960 and 2009 over the permafrost region even though model analyses indicate that warming alone would decrease soil C storage. Increases in gross primary production (GPP) largely explain the simulated increases in vegetation and soil C. The sensitivity of GPP to increases in atmospheric CO2 was the dominant cause of increases in GPP across the models, but comparison of simulated GPP trends across the 1982–2009 period with that of a global GPP data set indicates that all of the models overestimate the trend in GPP. Disturbance also appears to be an important factor affecting C storage, as models that consider disturbance had lower increases in C storage than models that did not consider disturbance. To improve the modeling of C in the permafrost region, there is the need for the modeling community to standardize structural representation of permafrost and carbon dynamics among models that are used to evaluate the permafrost C feedback and for the modeling and observational communities to jointly develop data sets and methodologies to more effectively benchmark models.
McHugh TA and Schwartz E (2016) A watering manipulation in a semiarid grassland induced changes in fungal but not bacterial community composition. Pedobiologia 59(3): 121–127.
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Read PublicationMonsoon precipitation in the arid southwestern United States is an important driver of ecosystem productivity, delivering up to 50% of annual precipitation during the summer months. These sporadic rainfall events typify drying-rewetting cycles and impose a physiological stress on the soil microbial communities responsible for carbon and nutrient cycling. As one aspect of climate change is an intensification of the hydrologic cycle, understanding how soil microbial communities and the processes they mediate are impacted by moisture fluctuations is increasingly important. We performed a month-long watering manipulation in the field and characterized bacterial and fungal communities across five time points using high-throughput sequencing. Watering treatment had a significant impact on fungal community composition, and there was a trend toward decreased fungal diversity and OTU richness in watered plots. In contrast, no significant differences were observed in bacterial communities between watered and control plots nor among sampling times. These findings suggest that fungi are more sensitive than bacteria to changes in soil moisture.
Mekonnen ZA, Grant RF, Schwalm CR (2016) Contrasting changes in gross primary productivity of different regions of North America as affected by warming in recent decades. Agricultural and Forest Meteorology 218-219.
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Read PublicationEcosystem responses to the increasing warming in recent decades across North America (NA) are spatially heterogeneous and partly uncertain. Here we examined the spatial and temporal variability of warming across different eco-regions of NA using long-term (1979–2010) climate data (North America Regional Reanalysis (NARR)) with 3-hourly time-step and 0.25° × 0.25° spatial resolution and run a comprehensive mathematical process model, <em>ecosys</em> to study the impacts of this variability in warming on gross primary productivity (GPP). In a site scale test of model results, annual GPP modeled for pixels which corresponded to the locations of 20 eddy covariance flux towers correlated well (<em>R</em><sup>2</sup> = 0.76) with annual GPP derived from the towers in 2005. At continental scale, long-term annual average modeled GPP correlated well (geographically weighed regression <em>R</em><sup>2</sup> = 0.8) with MODIS GPP. GPP modeled in eastern temperate forests and most areas with lower mean annual air temperature (<em>T</em><sub>a</sub>), such as those in northern forests and Taiga, increased due to early spring and late autumn warming observed in NARR and these eco-regions contributed 92% of the increases in NA GPP over the last three decades. However, modeled GPP declined in most southwestern regions of NA (accounting >50% of the ecosystems with declining GPP), due to water stress from rising <em>T</em><sub>a</sub> and declining precipitation, implying that further warming and projected dryness in this region could further reduce NA carbon uptake. Overall, NA modeled GPP increased by 5.8% in the last 30 years, with a positive trend of +0.012 ± 0.01 Pg C yr<sup>−1</sup> and a range of −1.16 to +0.87 Pg C yr<sup>−1</sup> caused by interannual variability of GPP from the long-term (1980–2010) mean. This variability was the greatest in southwest of US and part of the Great Plains, which could be as a result of frequent El Niño–Southern Oscillation’ events that led to major droughts.
Metcalfe AN, Kennedy TA, Muehlbauer JD (2016) Phenology of the adult angel lichen moth (Cisthene angelus) in Grand Canyon, USA. Southwestern Naturalist 61(3): 233-240.
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Read PublicationWe investigated the phenology of adult angel lichen moths (<i>Cisthene angelus</i>) along a 364-km long segment of the Colorado River in Grand Canyon, Arizona, USA, using a unique data set of 2,437 light-trap samples collected by citizen scientists. We found that adults of <i>C. angelus</i> were bivoltine from 2012 to 2014. We quantified plasticity in wing lengths and sex ratios among the two generations and across a 545-m elevation gradient. We found that abundance, but not wing length, increased at lower elevations and that the two generations differed in size and sex distributions. Our results shed light on the life history and morphology of a common, but poorly known, species of moth endemic to the southwestern United States and Mexico.
Morrissey EM, Mau RL, Schwartz E, Caporaso JG, Dijkstra P, van Gestel N, Koch BJ, Liu CM, Hayer M, McHugh TA, Marks JC, Price LB, and Hungate BA (2016) Phylogenetic organization of bacterial activity. The ISME Journal 10(9): 2336-2340.
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Read PublicationPhylogeny is an ecologically meaningful way to classify plants and animals, as closely related taxa frequently have similar ecological characteristics, functional traits and effects on ecosystem processes. For bacteria, however, phylogeny has been argued to be an unreliable indicator of an organism’s ecology owing to evolutionary processes more common to microbes such as gene loss and lateral gene transfer, as well as convergent evolution. Here we use advanced stable isotope probing with 13C and 18O to show that evolutionary history has ecological significance for in situ bacterial activity. Phylogenetic organization in the activity of bacteria sets the stage for characterizing the functional attributes of bacterial taxonomic groups. Connecting identity with function in this way will allow scientists to begin building a mechanistic understanding of how bacterial community composition regulates critical ecosystem functions.
Oberle B, Ogle K, Penagos JC, Sweeney J, Zanne A (2016) A Bayesian model for xylem vessel length accommodates subsampling and reveals skewed distributions in species that dominate seasonal habitats.. Journal of Plant Hydraulics 3: e-003, 17 pages.
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Read PublicationVessel length is an important but understudied dimension of variation in angiosperm vascular anatomy. Among other traits, vessel length mediates an important tradeoff between hydraulic efficiency and safety that could influence how plants respond to extreme weather with climate change. However, the functional significance of vessel length variation within individual stems is poorly known, in part because existing data analysis methods handle uncertainty in a way that makes vessel length distributions difficult to compare. We provide a solution to this problem through a hierarchical Bayesian framework for estimating vessel lengths and we demonstrate the flexibility of this method by applying it to data from serial cross sections of dye injected stems. Our approach can accelerate data collection and accommodate associated uncertainties by statistically correcting for bias and error that result from subsampling images. We illustrate our analytical framework by estimating and comparing vessel length distributions for 21 woody species characteristic of a North American forest. The best-fit model corrected for both bias due to secondary growth and sampling error within and among species. Vessel length estimates from this model varied by almost an order of magnitude and parameters of these distributions correlated with point estimates derived from a different, commonly used method. Furthermore, we show how key contrasts can be estimated with the Bayesian framework, and in doing so, we show that the shape of the vessel length distribution differed between ring- and diffuse-porous species, suggesting that within-stem vessel length variation corresponds to water stress seasonality and contributes to landscape-level habitat segregation. Our analysis method revealed the importance of within-stem variation in vessel length, and our results complement work on between-species variation in average vessel length, further illuminating how vascular anatomy can influence woody plants’ responses to water stress.
Ogle K, Barber JJ (2016) Plant and ecosystem memory. CHANCE 29(2): 16-22.
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Read PublicationOf course plants do not have brains and, thus, cannot actually remember what happened to them in the past. Although plants cannot remember, however, we use “memory” as a metaphor to refer to the effect of the past on current and future plant and ecosystem functioning. Such memory effects have been repeatedly shown in ecosystems (such as deserts) that are often defined by highly variable environmental conditions, where air temperatures, humidity, and soil water availability can differ greatly from one day to the next, week to week, among seasons, and year to year.
Ogle K, Ryan E, Dijkstra F, Pendall E (2016) Quantifying and reducing uncertainties in estimated soil CO2 fluxes with hierarchical data-model integration. Journal of Geophysical Research: Biogeosciences 121.
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Read PublicationNonsteady state chambers are often employed to measure soil CO<sub>2</sub> fluxes. CO<sub>2</sub> concentrations (<em>C</em>) in the headspace are sampled at different times (<em>t</em>), and fluxes (<em>f</em>) are calculated from regressions of <em>C</em> versus <em>t</em> based on a limited number of observations. Variability in the data can lead to poor fits and unreliable <em>f</em> estimates; groups with too few observations or poor fits are often discarded, resulting in “missing” <em>f</em> values. We solve these problems by fitting linear (steady state) and nonlinear (nonsteady state, diffusion based) models of <em>C</em> versus <em>t</em>, within a hierarchical Bayesian framework. Data are from the Prairie Heating and CO<sub>2</sub> Enrichment study that manipulated atmospheric CO<sub>2</sub>, temperature, soil moisture, and vegetation. CO<sub>2</sub> was collected from static chambers biweekly during five growing seasons, resulting in >12,000 samples and >3100 groups and associated fluxes. We compare <em>f</em> estimates based on nonhierarchical and hierarchical Bayesian (B versus HB) versions of the linear and diffusion-based (L versus D) models, resulting in four different models (BL, BD, HBL, and HBD). Three models fit the data exceptionally well (<em>R</em><sup>2</sup> ≥ 0.98), but the BD model was inferior (<em>R</em><sup>2</sup> = 0.87). The nonhierarchical models (BL and BD) produced highly uncertain <em>f</em> estimates (wide 95% credible intervals), whereas the hierarchical models (HBL and HBD) produced very precise estimates. Of the hierarchical versions, the linear model (HBL) underestimated <em>f</em> by ~33% relative to the nonsteady state model (HBD). The hierarchical models offer improvements upon traditional nonhierarchical approaches to estimating <em>f</em>, and we provide example code for the models.
Olefeldt D, Goswami S, Grosse G, Hayes D, Hugelius G, Kuhry P, McGuire AD, Romanovsky VE, Sannel ABK, Schuur EAG, Turetsky MR (2016) Circumpolar distribution and carbon storage of thermokarst landscapes. Nature Communications 7: 13043.
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Read PublicationThermokarst is the process whereby the thawing of ice-rich permafrost ground causes land subsidence, resulting in development of distinctive landforms. Accelerated thermokarst due to climate change will damage infrastructure, but also impact hydrology, ecology and biogeochemistry. Here, we present a circumpolar assessment of the distribution of thermokarst landscapes, defined as landscapes comprised of current thermokarst landforms and areas susceptible to future thermokarst development. At 3.6 × 10<sup>6</sup> km<sup>2</sup>, thermokarst landscapes are estimated to cover <span class="stix">∼</span>20% of the northern permafrost region, with approximately equal contributions from three landscape types where characteristic wetland, lake and hillslope thermokarst landforms occur. We estimate that approximately half of the below-ground organic carbon within the study region is stored in thermokarst landscapes. Our results highlight the importance of explicitly considering thermokarst when assessing impacts of climate change, including future landscape greenhouse gas emissions, and provide a means for assessing such impacts at the circumpolar scale.
Parker TC, Sadowsky J, Dunleavy H, Subke JA, Frey SD, Wookey PA (2016) Slowed Biogeochemical Cycling in Sub-arctic Birch Forest Linked to Reduced Mycorrhizal Growth and Community Change after a Defoliation Event. Ecosystems 1-15.
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Read PublicationSub-arctic birch forests (<em class="EmphasisTypeItalic ">Betula pubescens</em> Ehrh. ssp. <em class="EmphasisTypeItalic ">czerepanovii</em>) periodically suffer large-scale defoliation events caused by the caterpillars of the geometrid moths <em class="EmphasisTypeItalic ">Epirrita autumnata</em> and <em class="EmphasisTypeItalic ">Operophtera brumata</em>. Despite their obvious influence on ecosystem primary productivity, little is known about how the associated reduction in belowground C allocation affects soil processes. We quantified the soil response following a natural defoliation event in sub-arctic Sweden by measuring soil respiration, nitrogen availability and ectomycorrhizal fungi (EMF) hyphal production and root tip community composition. There was a reduction in soil respiration and an accumulation of soil inorganic N in defoliated plots, symptomatic of a slowdown of soil processes. This coincided with a reduction of EMF hyphal production and a shift in the EMF community to lower autotrophic C-demanding lineages (for example, /russula-lactarius). We show that microbial and nutrient cycling processes shift to a slower, less C-demanding state in response to canopy defoliation. We speculate that, amongst other factors, a reduction in the potential of EMF biomass to immobilise excess mineral nitrogen resulted in its build-up in the soil. These defoliation events are becoming more geographically widespread with climate warming, and could result in a fundamental shift in sub-arctic ecosystem processes and properties. EMF fungi may be important in mediating the response of soil cycles to defoliation and their role merits further investigation.
Peltier D, Fell M, Ogle K (2016) Legacy effects of drought in the southwestern United States: A multi-species synthesis. Ecological Monographs 86(3): 312-326.
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Read PublicationUnderstanding impacts of drought on tree growth and forest health is of major concern given projected climate change. Droughts may become more common in the Southwest due to extreme temperatures that will drive increased evapotranspiration and lower soil moisture, in combination with uncertain precipitation changes. Utilizing ~1.3 million tree-ring widths from the International Tree Ring Data Bank representing 10 species (eight conifers, two oaks) in the Southwest, we evaluated the effects of drought on tree growth. We categorized ring widths by formation year in relation to drought (pre-drought, drought year, and post-drought), and we used a mixed-effects model to estimate the effects of current and antecedent precipitation and temperature on tree growth during the post-drought recovery period. This allowed us to assess changes in sensitivity of tree growth to precipitation and temperature at multiple timescales following multiple droughts, and to evaluate drought resistance and recovery in these species. The effects of precipitation and temperature on ring widths following drought varied among species and time since drought. Across species, 16% of the climate effects (i.e., “sensitivities”) were significantly different from their pre-drought values. Species differed, with some showing increased sensitivities to precipitation and temperature following drought, and others showing decreased sensitivities. Furthermore, some species (e.g., <em>Abies concolor</em> and <em>Pinus ponderosa</em>) showed low resistance and slow recovery, with changes in growth sensitivities persisting up to 5 yr; others (e.g., <em>Juniper</em> spp.) showed high resistance, such that their climatic sensitivities did not change. Among species, the importance of different antecedent climate variables changed with time since drought. Though a majority of species responded positively to same-year precipitation pre-drought, all 10 species were positively affected by same-year precipitation the second year after drought. Our results demonstrate tree growth sensitivities vary among species and with time since drought, raising questions about physiological mechanisms and implications for forest health under future drought.
Salmon VG, Soucy P, Mauritz M, Celis G, Natali SM, Mack MC, Schuur EAG (2016) Nitrogen availability increases in a tundra ecosystem during five years of experimental permafrost thaw. Global Change Biology 22 (5): 1927-1941.
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Read PublicationPerennially frozen soil in high latitude ecosystems (permafrost) currently stores 1330-1580 Pg of carbon (C). As these ecosystems warm, the thaw and decomposition of permafrost is expected to release large amounts of C to the atmosphere. Fortunately, losses from the permafrost C pool will be partially offset by increased plant productivity. The degree to which plants are able to sequester C, however, will be determined by changing nitrogen (N) availability in these thawing soil profiles. N availability currently limits plant productivity in tundra ecosystems but plant access to N is expected improve as decomposition increases in speed and extends to deeper soil horizons. In order to evaluate the relationship between permafrost thaw and N availability, we monitored N cycling during five years of experimentally induced permafrost thaw at the Carbon in Permafrost Experimental Heating Research project (CiPEHR). Inorganic N availability increased significantly in response to deeper thaw and greater soil moisture induced by Soil warming. This treatment also prompted a 23% increase in aboveground biomass and a 49% increase in foliar N pools. The sedge Eriophorum vaginatum responded most strongly to warming: this species explained 91% of the change in aboveground biomass during the five year period. Air warming had little impact when applied alone, but when applied in combination with Soil warming, growing season soil inorganic N availability was significantly reduced. These results demonstrate that there is a strong positive relationship between the depth of permafrost thaw and N availability in tundra ecosystems but that this relationship can be diminished by interactions between increased thaw, warmer air temperatures and higher levels of soil moisture. Within five years of permafrost thaw, plants actively incorporate newly available N into biomass but C storage in live vascular plant biomass is unlikely to be greater than losses from deep soil C pools. This article is protected by copyright. All rights reserved.
Schädel C, Bader MK-F, Schuur EAG, Biasi C, Bracho R, Capek P, De Baets S, Diáková K, Ernakovich J, Estop-Aragones C, Graham DE, Hartley IP, Iversen CM, Kane E, Knoblauch C, Lupascu M, Martikainen PJ, Natali SM, Norby RJ, O’Donnell JA, Chowdhury TR, Šantrucková H, Shaver G, Sloan VL, Treat CC, Turetsky MR, Waldrop MP, Wickland KP (2016) Potential carbon emissions dominated by carbon dioxide from thawed permafrost soils. Nature Climate Change 6(10): 950-953.
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Read PublicationIncreasing temperatures in northern high latitudes are causing permafrost to thaw, making large amounts of previously frozen organic matter vulnerable to microbial decomposition. Permafrost thaw also creates a fragmented landscape of drier and wetter soil conditions that determine the amount and form (carbon dioxide (CO<sub>2</sub>), or methane (CH<sub>4</sub>)) 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 unclear. 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<span class="mb"><span class="mb"> </span></span>°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 CH<sub>4</sub>, 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 CO<sub>2</sub> will strengthen the permafrost C feedback more than waterlogged systems releasing CO<sub>2</sub> and CH<sub>4</sub> for a given amount of C.
Schuur EAG, Hugelius G (2016) Terrestrial Carbon Cycle. Arctic Report Card 2016.
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Read PublicationThe Arctic continues to warm at a rate that is currently twice as fast as the global average (see essay on <em><a href="http://www.arctic.noaa.gov/Report-Card/Report-Card-2016/ArtMID/5022/ArticleID/271/Surface-Air-Temperature">Surface Air Temperature</a></em>). Warming is causing normally frozen ground (permafrost) to thaw, exposing significant quantities of organic soil carbon to decomposition by soil microbes (Romanovsky et al. 2010, Romanovsky et al. 2012). This <strong>permafrost carbon</strong> is the remnants of plants, animals, and microbes accumulated in frozen soil over hundreds to thousands of years (Schuur et al. 2008). The northern permafrost zone holds twice as much carbon as currently in the atmosphere (Schuur et al. 2015, Hugelius et al. 2014, Tarnocai et al. 2009, Zimov et al. 2006). Release of just a fraction of this frozen carbon pool, as the greenhouse gases carbon dioxide and methane, into the atmosphere would dramatically increase the rate of future global climate warming (Schuur et al. 2013).
This report details recent advances in quantifying the amount of organic carbon stored in permafrost zone soils and recent trends (1970-2010) in the exchange of carbon between tundra ecosystems and the atmosphere. These data are the most recent comprehensive data synthesis across individual sites.
Schwartz E , Hayer M, Hungate BA, Koch BJ, McHugh TA, Mercurio W, Morrissey EM, Soldanova K (2016) Stable isotope probing with 18O-water to investigate microbial growth and death in environmental samples. Current Opinion in Biotechnology 41: 14–18.
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Read PublicationGrowth and mortality of microorganisms have been characterized through DNA stable isotope probing (SIP) with <sup>18</sup>O-water in soils from a range of ecosystems. Conventional SIP has been improved by sequencing a marker gene in all fractions retrieved from an ultracentrifuge tube to produce taxon density curves, which allow estimating the atom percent isotope composition of each microbial taxon's genome. Very recent advances in SIP with <sup>18</sup>O-water include expansion of the technique to aquatic samples, investigations of microbial turnover in soil, and the first use of <sup>18</sup>O-water in RNA-SIP studies.
Selmants PC, Adair KL, Litton CM, Giardina CP, Schwartz E (2016) Increases in mean annual temperature do not alter soil bacterial community structure in tropical montane wet forests. Ecosphere 7(4): Article e01296.
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Read PublicationSoil bacteria play a key role in regulating terrestrial biogeochemical cycling and greenhouse gas fluxes across the soil-atmosphere continuum. Despite their importance to ecosystem functioning, we lack a general understanding of how bacterial communities respond to climate change, especially in relatively understudied ecosystems like tropical montane wet forests. We used a well-studied and highly constrained 5.2°C mean annual temperature (MAT) gradient in tropical montane wet forests on the Island of Hawaii to test the hypothesis that long-term, whole-ecosystem warming and the accompanying increase in belowground carbon flux increase the diversity and alter the composition of soil bacterial communities. Across this MAT gradient, dominant vegetation, substrate type and age, soil moisture, and disturbance history are constant, allowing us to effectively isolate the influence of rising MAT on soil bacterial community structure. Contrary to our hypothesis, we found that the richness, evenness, and phylogenetic diversity of the soil bacterial community remained remarkably stable with MAT and that MAT did not predict variation in bacterial community composition despite a substantial increase in belowground soil carbon fluxes across the gradient. Our results suggest that other factors that are constant across this gradient—such as soil pH, water availability and plant species composition—may be more important than warming in influencing soil bacterial community composition and diversity, at least within the temperature range studied here (~13–18°C MAT). Ours is the first study to demonstrate stability of soil bacterial community structure with rising MAT and increased belowground carbon flux in a tropical wet forest ecosystem. Moreover, our results add to growing evidence that the diversity and composition of soil bacterial communities dominated by Proteobacteria and Acidobacteria in low-pH forest soils may be insensitive to the direct effect of climate warming.
Shao J, Zhou X, Luo Y, Zhang G, Yan W, Li J, Li B, Dan L, Fisher JB, Gao Z, He Y, Huntzinger D, Jain AK, Mao J, Meng J, Michalak AM, Parazoo NC, Peng C, Poulter B, Schwalm CR, Shi X, Sun R, Tao F, Tian H, Wei Y, Zeng N, Zhu Q, Zhu W (2016) Uncertainty analysis of terrestrial net primary productivity and net biome productivity in China during 1901–2005. Journal of Geophysical Research: Biogeosciences 121(5): 1372-1393.
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Read PublicationDespite the importance of net primary productivity (NPP) and net biome productivity (NBP), estimates of NPP and NBP for China are highly uncertain. To investigate the main sources of uncertainty, we synthesized model estimates of NPP and NBP for China from published literature and the Multi-scale Synthesis and Terrestrial Model Intercomparison Project (MsTMIP). The literature-based results showed that total NPP and NBP in China were 3.35 ± 1.25 and 0.14 ± 0.094 Pg C yr<sup>−1</sup>, respectively. Classification and regression tree analysis based on literature data showed that model type was the primary source of the uncertainty, explaining 36% and 64% of the variance in NPP and NBP, respectively. Spatiotemporal scales, land cover conditions, inclusion of the N cycle, and effects of N addition also contributed to the overall uncertainty. Results based on the MsTMIP data suggested that model structures were overwhelmingly important (>90%) for the overall uncertainty compared to simulations with different combinations of time-varying global change factors. The interannual pattern of NPP was similar among diverse studies and increased by 0.012 Pg C yr<sup>−1</sup> during 1981–2000. In addition, high uncertainty in China's NPP occurred in areas with high productivity, whereas NBP showed the opposite pattern. Our results suggest that to significantly reduce uncertainty in estimated NPP and NBP, model structures should be substantially tested on the basis of empirical results. To this end, coordinated distributed experiments with multiple global change factors might be a practical approach that can validate specific structures of different models.
Terrer C, Vicca S, Hungate BA, Phillips RP, Prentice IC (2016) Mycorrhizal association as a primary control of the CO2 fertilization effect. Science 353 (6294): 72.
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Read PublicationPlants buffer increasing atmospheric carbon dioxide (CO<sub>2</sub>) concentrations through enhanced growth, but the question whether nitrogen availability constrains the magnitude of this ecosystem service remains unresolved. Synthesizing experiments from around the world, we show that CO<sub>2</sub> fertilization is best explained by a simple interaction between nitrogen availability and mycorrhizal association. Plant species that associate with ectomycorrhizal fungi show a strong biomass increase (30 ± 3%, <em>P</em> < 0.001) in response to elevated CO<sub>2</sub> regardless of nitrogen availability, whereas low nitrogen availability limits CO<sub>2</sub> fertilization (0 ± 5%, <em>P</em> = 0.946) in plants that associate with arbuscular mycorrhizal fungi. The incorporation of mycorrhizae in global carbon cycle models is feasible, and crucial if we are to accurately project ecosystem responses and feedbacks to climate change.
Thomas RT, Prentice IC, Graven H, Ciais P, Fisher JB, Hayes DJ, Maoyi H, Huntzinger DN, Ito A, Jain A, Mao J, Michalak AM, Peng S, Poulter B, Ricciuto DM, Shi X, Schwalm CR, Tian H, Zeng N (2016) Increased light-use efficiency in northern terrestrial ecosystems indicated by CO2 and greening observations. Geophysical Research Letters 43(21): 11,339-11,349.
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Read PublicationObservations show an increasing amplitude in the seasonal cycle of CO<sub>2</sub> (ASC) north of 45°N of 56 ± 9.8% over the last 50 years and an increase in vegetation greenness of 7.5–15% in high northern latitudes since the 1980s. However, the causes of these changes remain uncertain. Historical simulations from terrestrial biosphere models in the Multiscale Synthesis and Terrestrial Model Intercomparison Project are compared to the ASC and greenness observations, using the TM3 atmospheric transport model to translate surface fluxes into CO<sub>2</sub>concentrations. We find that the modeled change in ASC is too small but the mean greening trend is generally captured. Modeled increases in greenness are primarily driven by warming, whereas ASC changes are primarily driven by increasing CO<sub>2</sub>. We suggest that increases in ecosystem-scale light use efficiency (LUE) have contributed to the observed ASC increase but are underestimated by current models. We highlight potential mechanisms that could increase modeled LUE.
Tian H, Lu C, Ciais P, Michalak AM, Canadell JG, Saikawa E, Huntzinger DN, Gurney KR, Sitch S, Zhang B, Yang J, Bousquet P, Bruhwiler L, Chen G, Dlugokencky E, Friedlingstein P, Melillo J, Pan S, Poulter B, Prinn R, Saunois M, Schwalm CR, Wofsy SC (2016) The terrestrial biosphere as a net source of greenhouse gases to the atmosphere. Nature 531(7593): 225-228.
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Read PublicationThe terrestrial biosphere can release or absorb the greenhouse gases, carbon dioxide (CO<sub>2</sub>), methane (CH<sub>4</sub>) and nitrous oxide (N<sub>2</sub>O), and therefore has an important role in regulating atmospheric composition and climate<sup><a id="ref-link-1" title="Lovelock, J. E. & Margulis, L. Atmospheric homeostasis by and for the biosphere: the gaia hypothesis. Tellus A 26, http://dx.doi.org/10.3402/tellusa.v26i1-2.9731 (1974)" href="http://www.nature.com/nature/journal/v531/n7593/full/nature16946.html#ref1">1</a></sup>. Anthropogenic activities such as land-use change, agriculture and waste management have altered terrestrial biogenic greenhouse gas fluxes, and the resulting increases in methane and nitrous oxide emissions in particular can contribute to climate change<sup><a id="ref-link-2" title="Vitousek, P. M., Mooney, H. A., Lubchenco, J. & Melillo, J. M. Human domination of Earth’s ecosystems. Science 277, 494–499 (1997)" href="http://www.nature.com/nature/journal/v531/n7593/full/nature16946.html#ref2">2</a>, <a id="ref-link-3" title="Ciais, P. et al. in Climate Change 2013: The Physical Science Basis (eds Stocker, T. F. et al. ) Ch. 6 (Cambridge Univ. Press, 2013)" href="http://www.nature.com/nature/journal/v531/n7593/full/nature16946.html#ref3">3</a></sup>. The terrestrial biogenic fluxes of individual greenhouse gases have been studied extensively<sup><a id="ref-link-4" title="Le Quéré, C. et al. Global carbon budget 2013. Earth Syst. Sci. Data 6, 235–263 (2014)" href="http://www.nature.com/nature/journal/v531/n7593/full/nature16946.html#ref4">4</a>, <a id="ref-link-5" title="Kirschke, S. et al. Three decades of global methane sources and sinks. Nature Geosci . 6, 813–823 (2013)" href="http://www.nature.com/nature/journal/v531/n7593/full/nature16946.html#ref5">5</a>, <a id="ref-link-6" title="Davidson, E. A. & Kanter, D. Inventories and scenarios of nitrous oxide emissions. Environ. Res. Lett. 9, 105012 (2014)" href="http://www.nature.com/nature/journal/v531/n7593/full/nature16946.html#ref6">6</a></sup>, but the net biogenic greenhouse gas balance resulting from anthropogenic activities and its effect on the climate system remains uncertain. Here we use bottom-up (inventory, statistical extrapolation of local flux measurements, and process-based modelling) and top-down (atmospheric inversions) approaches to quantify the global net biogenic greenhouse gas balance between 1981 and 2010 resulting from anthropogenic activities and its effect on the climate system. We find that the cumulative warming capacity of concurrent biogenic methane and nitrous oxide emissions is a factor of about two larger than the cooling effect resulting from the global land carbon dioxide uptake from 2001 to 2010. This results in a net positive cumulative impact of the three greenhouse gases on the planetary energy budget, with a best estimate (in petagrams of CO<sub>2</sub> equivalent per year) of 3.9 ± 3.8 (top down) and 5.4 ± 4.8 (bottom up) based on the GWP100 metric (global warming potential on a 100-year time horizon). Our findings suggest that a reduction in agricultural methane and nitrous oxide emissions, particularly in Southern Asia, may help mitigate climate change.
Tramontana G., Jung M, Schwalm CR, Ichii K, Camps-Valls G, Ráduly B, Reichstein M, Arain MA, Cescatti A, Kiely G, Merbold L, Serrano-Ortiz P, Sickert S, Wolf S, Papale D (2016) Predicting carbon dioxide and energy fluxes across global FLUXNET sites with regression algorithms. Biogeosciences 13(14): 4291-4313.
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Read PublicationSpatio-temporal fields of land–atmosphere fluxes derived from data-driven models can complement simulations by process-based land surface models. While a number of strategies for empirical models with eddy-covariance flux data have been applied, a systematic intercomparison of these methods has been missing so far. In this study, we performed a cross-validation experiment for predicting carbon dioxide, latent heat, sensible heat and net radiation fluxes across different ecosystem types with 11 machine learning (ML) methods from four different classes (kernel methods, neural networks, tree methods, and regression splines). We applied two complementary setups: (1) 8-day average fluxes based on remotely sensed data and (2) daily mean fluxes based on meteorological data and a mean seasonal cycle of remotely sensed variables. The patterns of predictions from different ML and experimental setups were highly consistent. There were systematic differences in performance among the fluxes, with the following ascending order: net ecosystem exchange (R 2 < 0.5), ecosystem respiration (R 2 > 0.6), gross primary production (R 2> 0.7), latent heat (R 2 > 0.7), sensible heat (R 2 > 0.7), and net radiation (R 2 > 0.8). The ML methods predicted the across-site variability and the mean seasonal cycle of the observed fluxes very well (R 2 > 0.7), while the 8-day deviations from the mean seasonal cycle were not well predicted (R 2 < 0.5). Fluxes were better predicted at forested and temperate climate sites than at sites in extreme climates or less represented by training data (e.g., the tropics). The evaluated large ensemble of ML-based models will be the basis of new global flux products.
Tucker C, Tamang S, Pendall E, Ogle K (2016) Shallow snowpack inhibits soil respiration in sagebrush steppe through multiple biotic and abiotic mechanisms. Ecosphere 7(5): e01297.
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Read PublicationIn sagebrush steppe, snowpack may govern soil respiration through its effect on multiple abiotic and biotic factors. Across the Intermountain West of the United States, snowpack has been declining for decades and is projected to decline further over the next century, making the response of soil respiration to snowpack a potentially important factor in the ecosystem carbon cycle. In this study, we evaluated the direct and indirect roles of the snowpack in driving soil respiration in sagebrush steppe ecosystems by taking advantage of highway snowfences in Wyoming to manipulate snowpack. An important contribution of this study is the use of Bayesian modeling to quantify the effects of soil moisture and temperature on soil respiration across a wide range of conditions from frozen to hot and dry, while simultaneously accounting for biotic factors (e.g., vegetation cover, root density, and microbial biomass and substrate-use diversity) affected by snowpack. Elevated snow depth increased soil temperature (in the winter) and moisture (winter and spring), and was associated with reduced vegetation cover and microbial biomass carbon. Soil respiration showed an exponential increase with temperature, with a temperature sensitivity that decreased with increasing seasonal temperature (<em>Q</em><sub>10</sub> = 4.3 [winter], 2.3 [spring], and 1.7 [summer]); frozen soils were associated with unrealistic <em>Q</em><sub>10</sub> ≈ 7989 due to the liquid-to-ice transition of soil water. Soil respiration was sensitive to soil water content; predicted respiration under very dry conditions was less than 10% of respiration under moist conditions. While higher vegetation cover increased soil respiration, this was not due to increased root density, and may reflect differences in litter inputs. Microbial substrate-use diversity was negatively related to reference respiration (i.e., respiration rate at a reference temperature and optimal soil moisture), although the mechanism remains unclear. This study indicates that soil respiration is inhibited by shallow snowpack through multiple mechanisms; thus, future decreases in snowpack across the sagebrush steppe have the potential to reduce losses of soil C, potentially affecting regional carbon balance.
van Gestel NC, Dhungana N, Tissue DT, Zak JC (2016) Seasonal microbial and nutrient responses during a 5-year reduction in the daily temperature range of soil in a Chihuahuan Desert ecosystem. Oecologia 180(1): 265-277.
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Read PublicationHigh daily temperature range of soil (DTRsoil) negatively affects soil microbial biomass and activity, but its interaction with seasonal soil moisture in regulating ecosystem function remains unclear. For our 5-year field study in the Chihuahuan Desert, we suspended shade cloth 15 cm above the soil surface to reduce daytime temperature and increase nighttime soil temperature compared to unshaded plots, thereby reducing DTRsoil (by 5 ºC at 0.2 cm depth) without altering mean temperatures. Microbial biomass production was primarily regulated by seasonal precipitation with the magnitude of the response dependent on DTRsoil. Reduced DTRsoil more consistently increased microbial biomass nitrogen (MBN; +38 %) than microbial biomass carbon (MBC) with treatment responses being similar in spring and summer. Soil respiration depended primarily on soil moisture with responses to reduced DTRsoil evident only in wetter summer soils (+53 %) and not in dry spring soils. Reduced DTRsoil had no effect on concentrations of dissolved organic C, soil organic matter (SOM), nor soil inorganic N (extractable NO3 −–N + NH4 +–N). Higher MBN without changes in soil inorganic N suggests faster N cycling rates or alternate sources of N. If N cycling rates increased without a change to external N inputs (atmospheric N deposition or N fixation), then productivity in this desert system, which is N-poor and low in SOM, could be negatively impacted with continued decreases in daily temperature range. Thus, the future N balance in arid ecosystems, under conditions of lower DTR, seems linked to future precipitation regimes through N deposition and regulation of soil heat load dynamics.
Walker X, Mack M, Johnstone J (2016) Predicting Ecosystem Resilience to Fire from Tree Ring Analysis in Black Spruce Forests. Ecosystems.
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Read PublicationClimate change has increased the occurrence, severity, and impact of disturbances on forested ecosystems worldwide, resulting in a need to identify factors that contribute to an ecosystem’s resilience or capacity to recover from disturbance. Forest resilience to disturbance may decline with climate change if mature trees are able to persist under stressful environmental conditions that do not permit successful recruitment and survival after a disturbance. In this study, we used the change in proportional representation of black spruce pre- to post-fire as a surrogate for resilience. We explored links between patterns of resilience and tree ring signals of drought stress across topographic moisture gradients within the boreal forest. We sampled 72 recently (2004) burned stands of black spruce in interior Alaska (USA); the relative dominance of black spruce after fire ranged from almost no change (high resilience) to a 90% decrease (low resilience). Variance partitioning analysis indicated that resilience was related to site environmental characteristics and climate–growth responses, with no unique contribution of pre-fire stand composition. The largest shifts in post-fire species composition occurred in sites that experienced the compounding effects of pre-fire drought stress and shallow post-fire organic layer thickness. These sites were generally located at warmer and drier landscape positions, suggesting they are less resilient to disturbance than sites in cool and moist locations. Climate–growth responses can provide an estimate of stand environmental stress to climate change and as such are a valuable tool for predicting landscape variations in forest ecosystem resilience.
Webb EE, Schuur EAG, Natali SM, Oken KL, Bracho R, Krapek JP, Risk D, Nickerson NR (2016) Increased wintertime CO2 loss as a result of sustained tundra warming. Journal of Geophysical Research: Biogeosciences 121(2): 249-265.
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Read PublicationPermafrost soils currently store approximately 1672 Pg of carbon (C), but as high latitudes warm, this temperature-protected C reservoir will become vulnerable to higher rates of decomposition. In recent decades, air temperatures in the high latitudes have warmed more than any other region globally, particularly during the winter. Over the coming century, the arctic winter is also expected to experience the most warming of any region or season, yet it is notably understudied. Here we present nonsummer season (NSS) CO2 flux data from the Carbon in Permafrost Experimental Heating Research project, an ecosystem warming experiment of moist acidic tussock tundra in interior Alaska. Our goals were to quantify the relationship between environmental variables and winter CO2 production, account for subnivean photosynthesis and late fall plant C uptake in our estimate of NSS CO2 exchange, constrain NSS CO2 loss estimates using multiple methods of measuring winter CO2 flux, and quantify the effect of winter soil warming on total NSS CO2 balance. We measured CO2 flux using four methods: two chamber techniques (the snow pit method and one where a chamber is left under the snow for the entire season), eddy covariance, and soda lime adsorption, and found that NSS CO2 loss varied up to fourfold, depending on the method used. CO2 production was dependent on soil temperature and day of season but atmospheric pressure and air temperature were also important in explaining CO2 diffusion out of the soil. Warming stimulated both ecosystem respiration and productivity during the NSS and increased overall CO2 loss during this period by 14% (this effect varied by year, ranging from 7 to 24%). When combined with the summertime CO2 fluxes from the same site, our results suggest that this subarctic tundra ecosystem is shifting away from its historical function as a C sink to a C source.
Wymore AS, Liu CM, Hungate BA, Schwartz E, Price LB, Whitham TG, Marks JC (2016) The Influence of Time and Plant Species on the Composition of the Decomposing Bacterial Community in a Stream Ecosystem. Microbial Ecology 71(4): 825-834.
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Read PublicationFoliar chemistry influences leaf decomposition, but little is known about how litter chemistry affects the assemblage of bacterial communities during decomposition. Here we examined relationships between initial litter chemistry and the composition of the bacterial community in a stream ecosystem. We incubated replicated genotypes of <em class="EmphasisTypeItalic ">Populus fremontii</em> and <em class="EmphasisTypeItalic ">P. angustifolia</em> leaf litter that differ in percent tannin and lignin, then followed changes in bacterial community composition during 28 days of decomposition using 16S rRNA gene-based pyrosequencing. Using a nested experimental design, the majority of variation in bacterial community composition was explained by time (i.e., harvest day) (<em class="EmphasisTypeItalic ">R</em> <em class="EmphasisTypeItalic ">2</em> = 0.50). Plant species, nested within harvest date, explained a significant but smaller proportion of the variation (<em class="EmphasisTypeItalic ">R</em> <em class="EmphasisTypeItalic ">2</em> = 0.03). Significant differences in community composition between leaf species were apparent at day 14, but no significant differences existed among genotypes. Foliar chemistry correlated significantly with community composition at day 14 (<em class="EmphasisTypeItalic ">r</em> = 0.46) indicating that leaf litter with more similar phytochemistry harbor bacterial communities that are alike. Bacteroidetes and β-proteobacteria dominated the bacterial assemblage on decomposing leaves, and Verrucomicrobia and α- and δ-proteobacteria became more abundant over time. After 14 days, bacterial diversity diverged significantly between leaf litter types with fast-decomposing <em class="EmphasisTypeItalic ">P. fremontii</em> hosting greater richness than slowly decomposing <em class="EmphasisTypeItalic ">P. angustifolia</em>; however, differences were no longer present after 28 days in the stream. Leaf litter tannin, lignin, and lignin: N ratios all correlated negatively with diversity. This work shows that the bacterial community on decomposing leaves in streams changes rapidly over time, influenced by leaf species via differences in genotype-level foliar chemistry.
Xue K, M Yuan M, J Shi Z, Qin Y, Deng Y, Cheng L, Wu L, He Z, Van Nostrand JD, Bracho R, Natali S, Schuur EAG, Luo C, Konstantinidis KT, Wang Q, Cole JR, Tiedje JM, Luo Y, Zhou J (2016) Tundra soil carbon is vulnerable to rapid microbial decomposition under climate warming. Nature Climate Change 6(6): 595-600.
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Read PublicationMicrobial decomposition of soil carbon in high-latitude tundra underlain with permafrost is one of the most important, but poorly understood, potential positive feedbacks of greenhouse gas emissions from terrestrial ecosystems into the atmosphere in a warmer world<sup><a title="Schuur, E. A. G. et al. Vulnerability of permafrost carbon to climate change: implications for the global carbon cycle. BioScience 58, 701-714 (2008)." href="http://www.nature.com/nclimate/journal/vaop/ncurrent/full/nclimate2940.html#ref1">1</a>, <a id="ref-link-5" title="Schuur, E. et al. Expert assessment of vulnerability of permafrost carbon to climate change. Climatic Change 119, 359-374 (2013)." href="http://www.nature.com/nclimate/journal/vaop/ncurrent/full/nclimate2940.html#ref2">2</a>, <a id="ref-link-6" title="Zhou, J. et al. Microbial mediation of carbon-cycle feedbacks to climate warming. Nature Clim. Change 2, 106-110 (2012)." href="http://www.nature.com/nclimate/journal/vaop/ncurrent/full/nclimate2940.html#ref3">3</a>, <a id="ref-link-7" title="Graham, D. E. et al. Microbes in thawing permafrost: the unknown variable in the climate change equation. ISME J. 6, 709-712 (2012)." href="http://www.nature.com/nclimate/journal/vaop/ncurrent/full/nclimate2940.html#ref4">4</a></sup>. Using integrated metagenomic technologies, we showed that the microbial functional community structure in the active layer of tundra soil was significantly altered after only 1.5 years of warming, a rapid response demonstrating the high sensitivity of this ecosystem to warming. The abundances of microbial functional genes involved in both aerobic and anaerobic carbon decomposition were also markedly increased by this short-term warming. Consistent with this, ecosystem respiration (<i>R</i><sub>eco</sub>) increased up to 38%. In addition, warming enhanced genes involved in nutrient cycling, which very likely contributed to an observed increase (30%) in gross primary productivity (GPP). However, the GPP increase did not offset the extra <i>R</i><sub>eco</sub>, resulting in significantly more net carbon loss in warmed plots compared with control plots. Altogether, our results demonstrate the vulnerability of active-layer soil carbon in this permafrost-based tundra ecosystem to climate warming and the importance of microbial communities in mediating such vulnerability.
Zscheischler J, Mahecha MD, Avitabile V, Calle L, Carvalhais N, Ciais P, Gans F, Gruber N, Hartmann J, Herold M, Ichii K, Jung M, Landschützer P, Laruelle GG, Lauerwald R, Papale D, Peylin P, Poulter B, Ray D, Regnier P, Rödenbeck C, Roman-Cuesta RM, Schwalm CR, Tramontana G, Tyukavina AT, Valentini R, van der Werf G, West TO, Wolf JE, Reichstein M (2016) An empirical spatiotemporal description of the global surface-atmosphere carbon fluxes: opportunities and data limitations. Biogeosciences Discuss. 1-32.
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Read PublicationUnderstanding the global carbon (C) cycle is of crucial importance to map current and future climate dynamics relative to global environmental change. A full characterization of C cycling requires detailed information on spatiotemporal patterns of surface-atmosphere fluxes. However, relevant C cycle observations are highly variable in their coverage and reporting standards. Especially problematic is the lack of integration of vertical oceanic, inland freshwaters and terrestrial carbon dioxide (CO<sub>2</sub>) exchange. Here we adopt a data-driven approach to synthesize a wide range of observation-based spatially explicit surface-atmosphere CO<sub>2</sub> fluxes from 2001 and 2010, to identify the state of today’s observational opportunities and data limitation. The considered fluxes include vertical net exchange of open oceans, continental shelves, estuaries, rivers, and lakes, as well as CO<sub>2</sub> fluxes related to gross primary productivity, terrestrial ecosystem respiration, fire emissions, loss of tropical aboveground C, harvested wood and crops, as well as fossil fuel and cement emissions. Spatially explicit CO<sub>2</sub> fluxes are obtained through geostatistical and/or remote sensing-based upscaling; minimizing biophysical or biogeochemical assumptions encoded in process-based models. We estimate a global bottom-up net C exchange (NCE) between the surface (land, ocean, and coastal areas) and the atmosphere. Uncertainties for NCE and its components are derived using resampling. In most continental regions our NCE estimates agree well with independent estimates from other sources. This holds for Europe (mean ±1 SD: 0.80 ± 0.16 PgC/yr, positive numbers are sources to the atmosphere), Russia (−0.02 ± 0.49 PgC/yr), East Asia (1.76 ± 0.38 PgC/yr), South Asia (0.25 ± 0.16 PgC/yr), and Australia (0.22 ± 0.47 PgC/yr). Our NCE estimates also suggest large C sink in tropical areas. The global NCE estimate is −6.07 ± 3.38 PgC/yr. This global bottom-up value is the opposite direction of what is expected from the atmospheric growth rate of CO<sub>2</sub>, and would require an offsetting surface C source of 4.27±0.10 PgC/yr. This mismatch highlights large knowledge and observational gaps in tropical areas, particularly in South America, Africa, and Southeast Asia, but also in North America. Our uncertainty assessment provides the basis for designing new observation campaigns. In particular, we lack seasonal monitoring of shelf, estuary and inland water-atmosphere C exchange. Also, extensive pCO<sub>2</sub>measurements are missing in the Southern Ocean. Most importantly, tropical land C fluxes suffer from a lack of in-situ observations. The consistent derivation of data uncertainties could serve as prior knowledge in multi-criteria optimization such as the Carbon Cycle Data Assimilation System (CCDAS) without overstating data credibility. Furthermore, the spatially explicit flux estimates may be used as a starting point to assess the validity of countries’ claims of reducing net C emissions in climate change negotiations.
2015
Adams KJ, Harrop B, James MA, and Marks JC (2015) Riparian Vegetation and Water Quality Monitoring: Middle Fossil Creek Riparian Habitat Protection and Restoration Project Final Report: Monitoring Data and Analysis, 2010-2014 (Task 4) Revised Final. Prepared for Coconino National Forest, Grant No. 09-162WPF, Arizona Water Protection Fund, July 17, 2015 2015.
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Download .PDFThe objective of the Fossil Creek Riparian Vegetation, Water Quality and Visitor Use
Monitoring Plan (NAU 2010) is to determine the effectiveness of the Middle Fossil Creek
Riparian Habitat Protection and Restoration Project. This final report discusses the results of
riparian vegetation and water quality monitoring conducted twice each year starting in 2010
(baseline monitoring) and for the following three and one-half years (2011, 2012, 2013, spring
2014). It also outlines the results of the recent spring 2014 monitoring effort.
Anderegg WRL, Schwalm C, Biondi F, Camarero JJ, Koch G, Litvak M, Ogle K, Shaw JD, Shevliakova E, Williams AP, Wolf A, Ziaco E, Pacala S. (2015) Pervasive drought legacies in forest ecosystems and their implications for carbon cycle models. Science 349 (6247): 528.
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Read PublicationThe impacts of climate extremes on terrestrial ecosystems are poorly understood but important for predicting carbon cycle feedbacks to climate change. Coupled climate–carbon cycle models typically assume that vegetation recovery from extreme drought is immediate and complete, which conflicts with the understanding of basic plant physiology. We examined the recovery of stem growth in trees after severe drought at 1338 forest sites across the globe, comprising 49,339 site-years, and compared the results with simulated recovery in climate-vegetation models. We found pervasive and substantial “legacy effects” of reduced growth and incomplete recovery for 1 to 4 years after severe drought. Legacy effects were most prevalent in dry ecosystems, among Pinaceae, and among species with low hydraulic safety margins. In contrast, limited or no legacy effects after drought were simulated by current climate-vegetation models. Our results highlight hysteresis in ecosystem-level carbon cycling and delayed recovery from climate extremes.
Barber JJ, Gupta P, Edwards W, Ogle K, Waller L (2015) Combining and comparing multiple serial dilution assays of particles in solution: application to brucellosis in elk of the Greater Yellowstone Ecosystem. Ecological & Environmental Statistics 22(1): 161-177.
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Read PublicationThe concentration detection threshold (CDT) is the concentration of particles in solution beyond which a (serial dilution) assay detects particle presence. By our account, CDTs typically are not estimated but are fixed at some value. Setting a CDT to zero (<span id="IEq1" class="InlineEquation"><span id="MathJax-Element-1-Frame" class="MathJax" style="box-sizing: inherit; display: inline-table; font-style: normal; font-weight: normal; line-height: normal; font-size: 12.75px; text-indent: 0px; text-align: left; text-transform: none; letter-spacing: normal; word-spacing: normal; word-wrap: normal; white-space: nowrap; float: none; direction: ltr; max-width: none; max-height: none; min-width: 0px; min-height: 0px; border: 0px; padding: 0px; margin: 0px; position: relative;" tabindex="0" data-mathml="<math xmlns="http://www.w3.org/1998/Math/MathML"><mi>d</mi><mo>=</mo><mn>0</mn></math>"><span id="MathJax-Span-1" class="math"><span id="MathJax-Span-2" class="mrow"><span id="MathJax-Span-3" class="mi">d</span><span id="MathJax-Span-4" class="mo">=</span><span id="MathJax-Span-5" class="mn">0</span></span></span><span class="MJX_Assistive_MathML">d=0</span></span></span>) implies perfect detection, a common assumption, and setting <span id="IEq2" class="InlineEquation"><span id="MathJax-Element-2-Frame" class="MathJax" style="box-sizing: inherit; display: inline-table; font-style: normal; font-weight: normal; line-height: normal; font-size: 12.75px; text-indent: 0px; text-align: left; text-transform: none; letter-spacing: normal; word-spacing: normal; word-wrap: normal; white-space: nowrap; float: none; direction: ltr; max-width: none; max-height: none; min-width: 0px; min-height: 0px; border: 0px; padding: 0px; margin: 0px; position: relative;" tabindex="0" data-mathml="<math xmlns="http://www.w3.org/1998/Math/MathML"><mi>d</mi><mo>&gt;</mo><mn>0</mn></math>"><span id="MathJax-Span-6" class="math"><span id="MathJax-Span-7" class="mrow"><span id="MathJax-Span-8" class="mi">d</span><span id="MathJax-Span-9" class="mo">></span><span id="MathJax-Span-10" class="mn">0</span></span></span><span class="MJX_Assistive_MathML">d>0</span></span></span> gives results that are “denominated” in units of <span id="IEq3" class="InlineEquation"><span id="MathJax-Element-3-Frame" class="MathJax" style="box-sizing: inherit; display: inline-table; font-style: normal; font-weight: normal; line-height: normal; font-size: 12.75px; text-indent: 0px; text-align: left; text-transform: none; letter-spacing: normal; word-spacing: normal; word-wrap: normal; white-space: nowrap; float: none; direction: ltr; max-width: none; max-height: none; min-width: 0px; min-height: 0px; border: 0px; padding: 0px; margin: 0px; position: relative;" tabindex="0" data-mathml="<math xmlns="http://www.w3.org/1998/Math/MathML"><mi>d</mi></math>"><span id="MathJax-Span-11" class="math"><span id="MathJax-Span-12" class="mrow"><span id="MathJax-Span-13" class="mi">d</span></span></span><span class="MJX_Assistive_MathML">d</span></span></span>, i.e., are relative to the choice of <span id="IEq4" class="InlineEquation"><span id="MathJax-Element-4-Frame" class="MathJax" style="box-sizing: inherit; display: inline-table; font-style: normal; font-weight: normal; line-height: normal; font-size: 12.75px; text-indent: 0px; text-align: left; text-transform: none; letter-spacing: normal; word-spacing: normal; word-wrap: normal; white-space: nowrap; float: none; direction: ltr; max-width: none; max-height: none; min-width: 0px; min-height: 0px; border: 0px; padding: 0px; margin: 0px; position: relative;" tabindex="0" data-mathml="<math xmlns="http://www.w3.org/1998/Math/MathML"><mi>d</mi></math>"><span id="MathJax-Span-14" class="math"><span id="MathJax-Span-15" class="mrow"><span id="MathJax-Span-16" class="mi">d</span></span></span><span class="MJX_Assistive_MathML">d</span></span></span>. Using multiple, different serial dilution assays, each with its own CDT, we choose a “reference assay,” to which we assign a fixed CDT value, to obtain relative estimates of the remaining assays’ CDTs and the underlying particle concentration. We present the CDTs as a novel way to account for or to compare different serial dilution assays, “sensitivities”. We apply our methodology to data from four assays of the presence of bacterial (<em class="EmphasisTypeItalic ">B. abortus</em>) antibodies in the serum of elk in the Greater Yellowstone Ecosystem, where transmission of brucellosis—the disease ensuing from infection—to commercial livestock is managed by the Wyoming Game and Fish Department to avoid the primary symptom of abnormal fetal abortion. Results agree qualitatively with the more traditional notion of sensitivity as the true positive rate.
Compson ZG, Hungate BA, Koch GW, Hart SC, Maestas JM, Adams KJ, Whitham TG, Marks JC (2015) Closely Related Tree Species Differentially Influence the Transfer of Carbon and Nitrogen from Leaf Litter Up the Aquatic Food Web. Ecosystems 18(2): 186-201.
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Read PublicationDecomposing leaf litter in streams provides habitat and nutrition for aquatic insects. Despite large differences in the nutritional qualities of litter among different plant species, their effects on aquatic insects are often difficult to detect. We evaluated how leaf litter of two dominant riparian species (Populus fremontii and P. angustifolia) influenced carbon and nitrogen assimilation by aquatic insect communities, quantifying assimilation rates using stable isotope tracers (13C, 15N). We tested the hypothesis that element fluxes from litter of different plant species better define aquatic insect community structure than insect relative abundances, which often fail. We found that (1) functional communities (defined by fluxes of carbon and nitrogen from leaf litter to insects) were different between leaf litter species, whereas more traditional insect communities (defined by relativized taxa abundances) were not different between leaf litter species, (2) insects assimilated N, but not C, at a higher rate from P. angustifolia litter compared to P. fremontii, even though P. angustifolia decomposes more slowly, and (3) the C:N ratio of material assimilated by aquatic insects was lower for P. angustifolia compared to P. fremontii, indicating higher nutritional quality, despite similar initial litter C:N ratios. These findings provide new evidence for the effects of terrestrial plant species on aquatic ecosystems via their direct influence on the transfer of elements up the food web. We demonstrate how isotopically labeled leaf litter can be used to assess the functioning of insect communities, uncovering patterns undetected by traditional approaches and improving our understanding of the association between food web structure and element cycling
Craine JM, Elmore AJ, Wang L, Augusto L, Baisden WT, Brookshire ENJ, Cramer MD, Hasselquist NJ, Hobbie EA, Kahmen A, Koba K, Kranabetter JM, Mack MC, Marin-Spiotta E, Mayor JR, Mclauchlan KK, Michelsen A, Nardoto GB, Oliveira RS, Perakis SS, Peri PL, Quesada CA, Richter A, Schipper LA, Stevenson BA, Turner BL, Viani RaG, Wanek W, Zeller B (2015) Convergence of soil nitrogen isotopes across global climate gradients. Scientific Reports 5: 8280.
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Read PublicationQuantifying global patterns of terrestrial nitrogen (N) cycling is central to predicting future patterns of primary productivity, carbon sequestration, nutrient fluxes to aquatic systems, and climate forcing. With limited direct measures of soil N cycling at the global scale, syntheses of the <sup>15</sup>N:<sup>14</sup>N ratio of soil organic matter across climate gradients provide key insights into understanding global patterns of N cycling. In synthesizing data from over 6000 soil samples, we show strong global relationships among soil N isotopes, mean annual temperature (MAT), mean annual precipitation (MAP), and the concentrations of organic carbon and clay in soil. In both hot ecosystems and dry ecosystems, soil organic matter was more enriched in <sup>15</sup>N than in corresponding cold ecosystems or wet ecosystems. Below a MAT of 9.8°C, soil δ<sup>15</sup>N was invariant with MAT. At the global scale, soil organic C concentrations also declined with increasing MAT and decreasing MAP. After standardizing for variation among mineral soils in soil C and clay concentrations, soil δ<sup>15</sup>N showed no consistent trends across global climate and latitudinal gradients. Our analyses could place new constraints on interpretations of patterns of ecosystem N cycling and global budgets of gaseous N loss.
Deane-Coe KK, Mauritz M, Celis G, Salmon V, Crummer KG, Natali SM, Schuur EAG (2015) Experimental Warming Alters Productivity and Isotopic Signatures of Tundra Mosses. Ecosystems 18 (6): 1070-1082.
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Read PublicationIn the tundra, mosses play an important functional role regulating belowground and ecosystem processes, but there is still considerable uncertainty about how tundra moss communities will respond to climate change. We examined the effects of 5 years of <em class="EmphasisTypeItalic ">in situ</em> air and soil warming on net primary productivity (NPP), carbon (C) and nitrogen (N) isotope signatures (δ<sup>13</sup>C and δ<sup>15</sup>N), and C:N in dominant Alaskan tundra mosses. Air warming increased mean air temperatures by up to 0.5°C and resulted in an 80–90% reduction in NPP in the feather moss <em class="EmphasisTypeItalic ">Pleurozium</em> and the peat moss <em class="EmphasisTypeItalic ">Sphagnum</em>. Soil warming increased permafrost thaw depth by 12–18%, upper soil water content by 23–27%, and resulted in a threefold increase in <em class="EmphasisTypeItalic ">Sphagnum</em> NPP. δ<sup>13</sup>C was positively correlated with moss NPP, and increased by 0.5–1‰ in all mosses under soil warming. C:N was reduced in <em class="EmphasisTypeItalic ">Sphagnum</em> and <em class="EmphasisTypeItalic ">Pleurozium</em>, due to increases in tissue %N in the soil warming treatment, suggesting that moss N availability could increase as temperatures increases. Higher N availability in warmer conditions, however, may be offset by unfavorable moisture conditions for moss growth. Similar to responses in tundra vascular plant communities, our results forecast interspecific differences in productivity among tundra mosses. Specifically, air warming may reduce productivity in <em class="EmphasisTypeItalic ">Sphagnum</em> and <em class="EmphasisTypeItalic ">Pleurozium</em>, but soil warming could offset this response in <em class="EmphasisTypeItalic ">Sphagnum</em>. Such responses may lead to changes in tundra moss community structure and function as temperatures increase that have the potential to alter tundra C and N cycling in a future climate.
Deng J, Gu Y, Zhang J, Xue K, Qin Y, Yuan M, Yin H, He Z, Wu L, Schuur EAG, Tiedje JM, Zhou J (2015) Shifts of tundra bacterial and archaeal communities along a permafrost thaw gradient in Alaska. Molecular Ecology 24(1): 222-234.
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Read PublicationUnderstanding the response of permafrost microbial communities to climate warming is crucial for evaluating ecosystem feedbacks to global change. This study investigated soil bacterial and archaeal communities by Illumina MiSeq sequencing of 16S rRNA gene amplicons across a permafrost thaw gradient at different depths in Alaska with thaw progression for over three decades. Over 4.6 million passing 16S rRNA gene sequences were obtained from a total of 97 samples, corresponding to 61 known classes and 470 genera. Soil depth and the associated soil physical–chemical properties had predominant impacts on the diversity and composition of the microbial communities. Both richness and evenness of the microbial communities decreased with soil depth. Acidobacteria, Verrucomicrobia, Alpha- and Gamma-Proteobacteria dominated the microbial communities in the upper horizon, whereas abundances of Bacteroidetes, Delta-Proteobacteria and Firmicutes increased towards deeper soils. Effects of thaw progression were absent in microbial communities in the near-surface organic soil, probably due to greater temperature variation. Thaw progression decreased the abundances of the majority of the associated taxa in the lower organic soil, but increased the abundances of those in the mineral soil, including groups potentially involved in recalcitrant C degradation (Actinomycetales, <em>Chitinophaga</em>, etc.). The changes in microbial communities may be related to altered soil C sources by thaw progression. Collectively, this study revealed different impacts of thaw in the organic and mineral horizons and suggests the importance of studying both the upper and deeper soils while evaluating microbial responses to permafrost thaw.
Dijkstra, Paul, Elena Salpas, Dawson Fairbanks, Erin B. Miller, Shannon B. Hagerty, Kees Jan van Groenigen, Bruce A. Hungate, Jane C. Marks, George W. Koch, Egbert Schwartz (2015) High carbon use efficiency in soil microbial communities is related to balanced growth, not storage compound synthesis. Soil Biology & Biochemistry (89) 35-43.
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Download .PDFThe efficiency with which microbes use substrate (Carbon Use Efficiency or CUE) to make new microbial biomass is an important variable in soil and ecosystem C cycling models. It is generally assumed that CUE of microbial activity in soils is low, however measured values vary widely. It is hypothesized that high values of CUE observed in especially short-term incubations reflect the build-up of storage compounds in response to a sudden increase in substrate availability and are therefore not representative of CUE of microbial activity in unamended soil. To test this hypothesis, we measured the 13CO2 release from six position-specific 13C-labeled glucose isotopomers in ponderosa pine and pinon-juniper soil. We compared this position-specific CO2 production pattern with patterns expected for 1) balanced microbial growth (synthesis of all compounds needed to build new microbial cells) at a low, medium, or high CUE, and 2) synthesis of storage compounds (glycogen, tri-palmitoyl-glycerol, and polyhydroxybutyrate). Results of this study show that synthesis of storage compounds is not responsible for the observed high CUE. Instead, it is the position-specific CO2 production expected for balanced growth and high CUE that best matches the observed CO2 production pattern in these two soils. Comparison with published studies suggests that the amount of glucose added in this study is too low and the duration of the experiment too short to affect microbial metabolism. We conclude that the hypothesis of high CUE in undisturbed soil microbial communities remains viable and worthy of further testing.
Duval BD, Natali SM, Hungate BA (2015) What Constitutes Plant-Available Molybdenum in Sandy Acidic Soils?. Communications in Soil Science and Plant Analysis 46 (3): 318-326.
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Read PublicationMolybdenum (Mo) is critical for the function of enzymes related to nitrogen cycling. Concentrations of Mo are very low in sandy, acidic soils, and biologically available Mo is only a small fraction of the total pool. While several methods have been proposed to measure plant-available Mo, there has not been a recent comprehensive analytical study that compares soil extraction methods as predictors of plant Mo uptake. A suite of five assays [total acid microwave digestion, ethylenediamenetetraaacetic acid (EDTA) extraction, Environmental Protection Agency (EPA) protocol 3050B, ammonium oxalate extraction, and pressurized hot water] was employed, followed by the determination of soil Mo concentrations via inductively coupled mass spectroscopy. The concentrations of soil Mo determined from these assays and their relationships as predictors of plant Mo concentration were compared. The assays yielded different concentrations of Mo: total digest > EPA > ammonium oxalate ≥ EDTA > pressurized hot water. Legume foliar Mo concentrations were most strongly correlated with ammonium oxalate–extractable Mo from soils, but an oak species showed no relationship with any soil Mo fraction and foliar Mo. Bulk fine roots in the 10- to 30-cm soil horizon were significantly correlated with the ammonium oxalate Mo fraction. There were significant correlations between ammonium oxalate Mo and the oxides of iron (Fe), manganese (Mn), and aluminum (Al). Results suggest that the ammonium oxalate extraction for soil Mo is the best predictor of plant-available Mo for species with high Mo requirements such as legumes and that plant-available Mo tracks strongly with other metal oxides in sandy, acidic soils.
Ha W, Kolb TE, Springer AE, Dore S, O'Donnell FC, Martinez Morales R, Masek Lopez S, Koch GW (2015) Evapotranspiration comparisons between eddy covariance measurements and meteorological and remote-sensing-based models in disturbed ponderosa pine forests. Ecohydrology 8(7): 1335-1350.
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Read PublicationEvapotranspiration (ET) comprises a major portion of the water budget in forests, yet few studies have measured or estimated ET in semi-arid, high-elevation ponderosa pine forests of the south-western USA or have investigated the capacity of models to predict ET in disturbed forests. We measured actual ET with the eddy covariance (eddy) method over 4 years in three ponderosa pine forests near Flagstaff, Arizona, that differ in disturbance history (undisturbed control, wildfire burned, and restoration thinning) and compared these measurements (415–510 mm year<sup>−1</sup> on average) with actual ET estimated from five meteorological models [Penman–Monteith (P-M), P-M with dynamic control of stomatal resistance (P-M-d), Priestley–Taylor (P-T), McNaughton–Black (M-B), and Shuttleworth–Wallace (S-W)] and from the Moderate Resolution Imaging Spectroradiometer (MODIS) ET product. The meteorological models with constant stomatal resistance (P-M, M-B, and S-W) provided the most accurate estimates of annual eddy ET (average percent differences ranged between 11 and −14%), but their accuracy varied across sites. The P-M-d consistently underpredicted ET at all sites. The more simplistic P-T model performed well at the control site (18% overprediction) but strongly overpredicted annual eddy ET at the restoration sites (92%) and underpredicted at the fire site (−26%). The MODIS ET underpredicted annual eddy ET at all sites by at least 51% primarily because of underestimation of leaf area index. Overall, we conclude that with accurate parameterization, micrometeorological models can predict ET within 30% in forests of the south-western USA and that remote sensing-based ET estimates need to be improved through use of higher resolution products.
Hicks Pries CE, Logtestijn RSP, Schuur EAG, Natali SM, Cornelissen JHC, Aerts R, Dorrepaal E (2015) Decadal warming causes a consistent and persistent shift from heterotrophic to autotrophic respiration in contrasting permafrost ecosystems. Global Change Biology 21 (12): 4508-4519.
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Read PublicationSoil carbon in permafrost ecosystems has the potential to become a major positive feedback to climate change if permafrost thaw increases heterotrophic decomposition. However, warming can also stimulate autotrophic production leading to increased ecosystem carbon storage—a negative climate change feedback. Few studies partitioning ecosystem respiration examine decadal warming effects or compare responses among ecosystems. Here, we first examined how 11 years of warming during different seasons affected autotrophic and heterotrophic respiration in a bryophyte-dominated peatland in Abisko, Sweden. We used natural abundance radiocarbon to partition ecosystem respiration into autotrophic respiration, associated with production, and heterotrophic decomposition. Summertime warming decreased the age of carbon respired by the ecosystem due to increased proportional contributions from autotrophic and young soil respiration and decreased proportional contributions from old soil. Summertime warming's large effect was due to not only warmer air temperatures during the growing season, but also to warmer deep soils year-round. Second, we compared ecosystem respiration responses between two contrasting ecosystems, the Abisko peatland and a tussock-dominated tundra in Healy, Alaska. Each ecosystem had two different timescales of warming (<5 years and over a decade). Despite the Abisko peatland having greater ecosystem respiration and larger contributions from heterotrophic respiration than the Healy tundra, both systems responded consistently to short- and long-term warming with increased respiration, increased autotrophic contributions to ecosystem respiration, and increased ratios of autotrophic to heterotrophic respiration. We did not detect an increase in old soil carbon losses with warming at either site. If increased autotrophic respiration is balanced by increased primary production, as is the case in the Healy tundra, warming will not cause these ecosystems to become growing season carbon sources. Warming instead causes a persistent shift from heterotrophic to more autotrophic control of the growing season carbon cycle in these carbon-rich permafrost ecosystems.
Hicks Pries CE, Schuur EAG, Natali SM, Crummer KG (2015) Old soil carbon losses increase with ecosystem respiration in experimentally thawed tundra. Nature Climate Change 6(2): 214–218.
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Read PublicationOld soil carbon (C) respired to the atmosphere as a result of permafrost thaw has the potential to become a large positive feedback to climate change. As permafrost thaws, quantifying old soil contributions to ecosystem respiration (<i>R</i><sub>eco</sub>) and understanding how these contributions change with warming is necessary to estimate the size of this positive feedback. We used naturally occurring C isotopes (δ<sup>13</sup>C and Δ<sup>14</sup>C) to partition <i>R</i><sub>eco</sub> into plant, young soil and old soil sources in a subarctic air and soil warming experiment over three years. We found that old soil contributions to <i>R</i><sub>eco</sub> increased with soil temperature and <i>R</i><sub>eco</sub> flux. However, the increase in the soil warming treatment was smaller than expected because experimentally warming the soils increased plant contributions to <i>R</i><sub>eco</sub> by 30%. On the basis of these data, an increase in mean annual temperature from −5 to 0<span class="mb"><span class="mb"> </span></span>°C will increase old soil C losses from moist acidic tundra by 35–55<span class="mb"><span class="mb"> </span></span>g C<span class="mb"><span class="mb"> </span></span>m<sup>−2</sup> during the growing season. The largest losses will probably occur where the plant response to warming is minimal.
Hungate BA, Mau RL, Schwartz E, Caporaso JG, Dijkstra P, van Gestel N, Koch BJ, Liu CM, McHugh TA, Marks JC, Morrissey EM, Price LB (2015) Quantitative Microbial Ecology Through Stable Isotope Probing. Applied and Environmental Microbiology 81(21): 7570-7581.
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Read PublicationBacteria grow and transform elements at different rates, and as yet, quantifying this variation in the environment is difficult. Determining isotope enrichment with fine taxonomic resolution after exposure to isotope tracers could help, but there are few suitable techniques. We propose a modification to stable isotope probing (SIP) that enables the isotopic composition of DNA from individual bacterial taxa after exposure to isotope tracers to be determined. In our modification, after isopycnic centrifugation, DNA is collected in multiple density fractions, and each fraction is sequenced separately. Taxon-specific density curves are produced for labeled and nonlabeled treatments, from which the shift in density for each individual taxon in response to isotope labeling is calculated. Expressing each taxon's density shift relative to that taxon's density measured without isotope enrichment accounts for the influence of nucleic acid composition on density and isolates the influence of isotope tracer assimilation. The shift in density translates quantitatively to isotopic enrichment. Because this revision to SIP allows quantitative measurements of isotope enrichment, we propose to call it quantitative stable isotope probing (qSIP). We demonstrated qSIP using soil incubations, in which soil bacteria exhibited strong taxonomic variations in 18O and 13C composition after exposure to [18O]water or [13C]glucose. The addition of glucose increased the assimilation of 18O into DNA from [18O]water. However, the increase in 18O assimilation was greater than expected based on utilization of glucose-derived carbon alone, because the addition of glucose indirectly stimulated bacteria to utilize other substrates for growth. This example illustrates the benefit of a quantitative approach to stable isotope probing.
Ji L, Wylie BK, Brown DRN, Peterson B, Alexander HD, Mack MC, Rover J, Waldrop MP, Mcfarland JW, Chen X, Pastick NJ (2015) Spatially explicit estimation of aboveground boreal forest biomass in the Yukon River Basin, Alaska. International Journal of Remote Sensing 36(4): 939-953.
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Read PublicationQuantification of aboveground biomass (AGB) in Alaska's boreal forest is essential to the accurate evaluation of terrestrial carbon stocks and dynamics in northern high-latitude ecosystems. Our goal was to map AGB at 30 m resolution for the boreal forest in the Yukon River Basin of Alaska using Landsat data and ground measurements. We acquired Landsat images to generate a 3-year (2008-2010) composite of top-of-atmosphere reflectance for six bands as well as the brightness temperature (BT). We constructed a multiple regression model using field-observed AGB and Landsat-derived reflectance, BT, and vegetation indices. A basin-wide boreal forest AGB map at 30 m resolution was generated by applying the regression model to the Landsat composite. The fivefold cross-validation with field measurements had a mean absolute error (MAE) of 25.7 Mg ha<sup>−1</sup> (relative MAE 47.5%) and a mean bias error (MBE) of 4.3 Mg ha<sup>−1</sup> (relative MBE 7.9%). The boreal forest AGB product was compared with lidar-based vegetation height data; the comparison indicated that there was a significant correlation between the two data sets.
Koch GW, Sillett SC, Antoine ME, Williams CB (2015) Growth maximization trumps maintenance of leaf conductance in the tallest angiosperm. Oecologia 177(2): 321-331.
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Read PublicationStructural and physiological changes that occur as trees grow taller are associated with increased hydraulic constraints on leaf gas exchange, yet it is unclear if leaf-level constraints influence whole-tree growth as trees approach their maximum size. We examined variation in leaf physiology, leaf area to sapwood area ratio (<em class="EmphasisTypeItalic ">L</em>/<em class="EmphasisTypeItalic ">S</em>), and annual aboveground growth across a range of tree heights in<em class="EmphasisTypeItalic "> Eucalyptus regnans</em>. Leaf photosynthetic capacity did not differ among upper crown leaves of individuals 61.1–92.4 m tall. Maximum daily and integrated diurnal stomatal conductance (<em class="EmphasisTypeItalic ">g</em><sub>s</sub>) averaged 36 and 34 % higher, respectively, in upper crown leaves of ~60-m-tall, 80-year-old trees than in ~90-m-tall, 300-year-old trees, with larger differences observed on days with a high vapor pressure deficit (VPD). Greater stomatal regulation in taller trees resulted in similar minimum daily leaf water potentials (<em class="EmphasisTypeItalic ">Ψ</em><sub>L</sub>) in shorter and taller trees over a broad range of VPDs. The long-term stomatal limitation on photosynthesis, as inferred from leaf <em class="EmphasisTypeItalic ">δ</em><sup>13</sup>C composition, was also greater in taller trees. The <em class="EmphasisTypeItalic ">δ</em><sup>13</sup>C of wood indicated that the bulk of photosynthesis used to fuel wood production in the main trunk and branches occurred in the upper crown. <em class="EmphasisTypeItalic ">L</em>/<em class="EmphasisTypeItalic ">S</em> increased with tree height, especially after accounting for size-independent variation in crown structure across 27 trees up to 99.8 m tall. Despite greater stomatal limitation of leaf photosynthesis in taller trees, total <em class="EmphasisTypeItalic ">L</em> explained 95 % of the variation in annual aboveground biomass growth among 15 trees measured for annual biomass growth increment in 2006. Our results support a theoretical model proposing that, in the face of increasing hydraulic constraints with height, whole-tree growth is maximized by a resource trade-off that increases <em class="EmphasisTypeItalic ">L</em> to maximize light capture rather than by reducing <em class="EmphasisTypeItalic ">L</em>/<em class="EmphasisTypeItalic ">S</em> to sustain <em class="EmphasisTypeItalic ">g</em><sub>s</sub>.
Koven CD, Schuur EAG, Schädel C, Bohn TJ, Burke EJ, Chen G, Chen X, Ciais P, Grosse G, Harden JW, Hayes DJ, Hugelius G, Jafarov EE, Krinner G, Kuhry P, Lawrence DM, Macdougall AH, Marchenko SS, Mcguire AD, Natali SM, Nicolsky DJ, Olefeldt D, Peng S, Romanovsky VE, Schaefer KM, Strauss J, Treat CC, Turetsky M (2015) A simplified, data-constrained approach to estimate the permafrost carbon–climate feedback. Philosophical Transactions of the Royal Society of London A: Mathematical, Physical and Engineering Sciences 373(2054).
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Read PublicationWe present an approach to estimate the feedback from large-scale thawing of permafrost soils using a simplified, data-constrained model that combines three elements: soil carbon (C) maps and profiles to identify the distribution and type of C in permafrost soils; incubation experiments to quantify the rates of C lost after thaw; and models of soil thermal dynamics in response to climate warming. We call the approach the Permafrost Carbon Network Incubation–Panarctic Thermal scaling approach (PInc-PanTher). The approach assumes that C stocks do not decompose at all when frozen, but once thawed follow set decomposition trajectories as a function of soil temperature. The trajectories are determined according to a three-pool decomposition model fitted to incubation data using parameters specific to soil horizon types. We calculate litterfall C inputs required to maintain steady-state C balance for the current climate, and hold those inputs constant. Soil temperatures are taken from the soil thermal modules of ecosystem model simulations forced by a common set of future climate change anomalies under two warming scenarios over the period 2010 to 2100. Under a medium warming scenario (RCP4.5), the approach projects permafrost soil C losses of 12.2–33.4 Pg C; under a high warming scenario (RCP8.5), the approach projects C losses of 27.9–112.6 Pg C. Projected C losses are roughly linearly proportional to global temperature changes across the two scenarios. These results indicate a global sensitivity of frozen soil C to climate change (<em>γ</em> sensitivity) of −14 to −19 Pg C °C<sup>−1</sup> on a 100 year time scale. For CH<sub>4</sub> emissions, our approach assumes a fixed saturated area and that increases in CH<sub>4</sub> emissions are related to increased heterotrophic respiration in anoxic soil, yielding CH<sub>4</sub> emission increases of 7% and 35% for the RCP4.5 and RCP8.5 scenarios, respectively, which add an additional greenhouse gas forcing of approximately 10–18%. The simplified approach presented here neglects many important processes that may amplify or mitigate C release from permafrost soils, but serves as a data-constrained estimate on the forced, large-scale permafrost C response to warming.
Kropp H, Ogle K (2015) Seasonal stomatal behavior of a common desert shrub and the influence of plant neighbors. Oecologia 177(2): 345-355.
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Read PublicationStomata simultaneously regulate plant carbon gain and water loss, and patterns of stomatal conductance (<em class="EmphasisTypeItalic ">g</em><sub>s</sub>) provide insight into water use strategies. In arid systems, <em class="EmphasisTypeItalic ">g</em><sub>s</sub> varies seasonally based on factors such as water availability and temperature. Moreover, the presence and species identity of neighboring plants likely affects <em class="EmphasisTypeItalic ">g</em><sub>s</sub> of the focal plant by altering available soil water and microclimate conditions. We investigated stomatal behavior in <em class="EmphasisTypeItalic ">Larrea tridentata</em>, a drought-tolerant, evergreen shrub occurring throughout the arid southwestern United States. We measured <em class="EmphasisTypeItalic ">g</em><sub>s</sub> in <em class="EmphasisTypeItalic ">Larrea</em> over multiple seasons in the presence of neighbors representing different woody species. The data were analyzed in the context of a commonly used phenomenological model that relates <em class="EmphasisTypeItalic ">g</em><sub>s</sub> to vapor pressure deficit (<em class="EmphasisTypeItalic ">D</em>) to understand spatial and temporal differences in stomatal behavior. We found that <em class="EmphasisTypeItalic ">g</em><sub>s</sub> in <em class="EmphasisTypeItalic ">Larrea</em> was affected by neighborhood association, and these effects varied seasonally. The greatest effect of neighborhood association on <em class="EmphasisTypeItalic ">g</em><sub>s</sub> occurred during the winter period, where <em class="EmphasisTypeItalic ">Larrea</em> growing alone (without neighbors) had higher <em class="EmphasisTypeItalic ">g</em><sub>s</sub> compared to <em class="EmphasisTypeItalic ">Larrea</em> growing with neighbors. <em class="EmphasisTypeItalic ">Larrea</em>’s stomatal sensitivity to <em class="EmphasisTypeItalic ">D</em> and reference conductance (i.e., <em class="EmphasisTypeItalic ">g</em><sub>s</sub> at <em class="EmphasisTypeItalic ">D</em> = 1 kPa) also differed significantly among different neighbor associations. Random effects indicated reference <em class="EmphasisTypeItalic ">g</em><sub>s</sub> varied over short time scales (daily), while stomatal sensitivity showed little daily or seasonal variation, but was notably affected by neighbor associations such that neighboring species, especially trees, reduced <em class="EmphasisTypeItalic ">Larrea</em>’s sensitivity to <em class="EmphasisTypeItalic ">D</em>. Overall, seasonal dynamics and neighborhood conditions appear critical to understanding temporal and spatial variation in <em class="EmphasisTypeItalic ">Larrea</em>’s physiological behavior.
Lamit LJ, Busby PE, Lau MK, Compson ZG, Wojtowicz T, Keith AR, Zinkgraf MS, Schweitzer JA, Shuster SM, Gehring CA, and Whitham TG (2015) Tree genotype mediates covariance among communities from microbes to lichens and arthropods. Journal of Ecology 103(4): 840–850.
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Read PublicationCommunity genetics studies frequently focus on individual communities associated with individual plant genotypes, but little is known about the genetically based relationships among taxonomically and spatially disparate communities. We integrate studies of a wide range of communities living on the same plant genotypes to understand how the ecological and evolutionary dynamics of one community may be constrained or modulated by its underlying genetic connections to another community. We use pre-existing data sets collected from Populus angustifolia (narrowleaf cottonwood) growing in a common garden to test the hypothesis that the composition of pairs of distinct communities (e.g. endophytes, pathogens, lichens, arthropods, soil microbes) covary across tree genotypes, such that individual plant genotypes that support a unique composition of one community are more likely to support a unique composition of another community. We then evaluate the hypotheses that physical proximity, taxonomic similarity, time between sampling (time attenuation), and interacting foundation species within communities explain the strength of correlations. Three main results emerged. First, Mantel tests between communities revealed moderate to strong (q = 0.25–0.85) community–genetic correlations in almost half of the comparisons; correlations among phyllosphere endophyte, pathogen and arthropod communities were the most robust. Secondly, physical proximity determined the strength of community–genetic correlations, supporting a physical proximity hypothesis. Thirdly, consistent with the interacting foundation species hypothesis, the most abundant species drove many of the stronger correlations. Other hypotheses were not supported. Synthesis. The field of community genetics demonstrates that the structure of communities varies among plant genotypes; our results add to this field by showing that disparate communities covary among plant genotypes. Eco-evolutionary dynamics between plants and their associated organisms may therefore be mediated by the shared connections of different communities to plant genotype, indicating that the organization of biodiversity in this system is genetically based and non-neutral.
LeRoy CJ, Fischer DG, Andrews WM, Belleveau L, Barlow CH, Schweitzer JA, Bailey JK, Marks JC, Kallestad JC (2015) Salmon carcasses influence genetic linkages between forests and streams. Canadian Journal of Fisheries and Aquatic Sciences 73(6): 910-920.
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Read PublicationBiodiversity at many scales (functional group, species, genetic) can result in emergent ecological patterns. Here we explore the influence of tree genotypic variation and diversity on in-stream ecosystem processes and aquatic communities. We test whether genetically diverse inputs of leaf litter interact with a keystone organism, anadromous salmon, to influence in-stream ecosystem function. We used reach-level manipulation of salmon carcasses and leaf litter bags to examine how nutrient inputs interact with genetic variation in leaf litter decomposition. Genotypic variation in black cottonwood (<i>Populus balsamifera</i> ssp. <i>trichocarpa</i>) significantly influenced leaf litter chemistry, litter mass loss, and fungal biomass, but these variables were only weakly influenced by salmon carcass presence or a genotype × salmon (G × E) interaction. Mixtures of genotypes tended to demonstrate antagonistic effects (slower than expected decomposition) in the absence of salmon, but synergistic effects (faster than expected decomposition) when salmon were present. Our findings suggest that the influence of plant genotypic variation in linking aquatic and terrestrial ecosystems may be altered and in some cases intensified in the presence of a keystone vertebrate species.
Liang J, Li D, Shi Z, Tiedje JM, Zhou J, Schuur EAG, Konstantinidis KT, Luo Y (2015) Methods for estimating temperature sensitivity of soil organic matter based on incubation data: A comparative evaluation. Soil Biology and Biochemistry.
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Read PublicationAlthough the temperature sensitivity (Q<sub>10</sub>) of soil organic matter (SOM) decomposition has been widely studied, the estimate substantially depends on the methods used with specific assumptions. Here we compared several commonly used methods (i.e., one-pool (1P) model, two-discrete-pool (2P) model, three-discrete-pool (3P) model, and time-for-substrate (T4S) Q<sub>10</sub> method) plus a new and more process-oriented approach for estimating Q<sub>10</sub> of SOM decomposition from laboratory incubation data to evaluate the influences of the different methods and assumptions on Q<sub>10</sub> estimation. The process-oriented approach is a three-transfer-pool (3PX) model that resembles the decomposition sub-model commonly used in Earth system models. The temperature sensitivity and other parameters in the models were estimated from the cumulative CO<sub>2</sub>emission using the Bayesian Markov Chain Monte Carlo (MCMC) technique. The estimated Q<sub>10</sub>s generally increased with the soil recalcitrance, but decreased with the incubation temperature increase. Our results indicated that the 1P model did not adequately simulate the dynamics of SOM decomposition and thus was not adequate for the Q<sub>10</sub> estimation. All the multi-pool models fitted the soil incubation data well. The Akaike information criterion (<em>AIC</em>) analysis suggested that the 2P model is the most parsimonious. As the incubation progressed, Q<sub>10</sub> estimated by the 3PX model was smaller than those by the 2P and 3P models because the continuous C transfers from the slow and passive pools to the active pool were included in the 3PX model. Although the T4S method could estimate the Q<sub>10</sub> of labile carbon appropriately, our analyses showed that it overestimated that of recalcitrant SOM. The similar structure of 3PX model with the decomposition sub-model of Earth system models provides a possible approach, via the data assimilation techniques, to incorporate results from numerous incubation experiments into Earth system models.
Liu CM, Hungate BA, Tobiana, AAR, Ravele R, Prodgerf JL, Serwaddag D, Kigozig G, Galiwangog RM, Nalugodag F, Keim P, Wawerh MJ, Price LB, Gray RH (2015) Penile Microbiota and Female Partner Bacterial Vaginosis in Rakai, Uganda. mBio 6 (3): e00589-15.
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Download .PDFBacterial vaginosis (BV) is a common vaginal bacterial imbalance associated with risk for HIV and poor gynecologic and obstetric outcomes. Male circumcision reduces BV-associated bacteria on the penis and decreases BV in female partners, but the link between penile microbiota and female partner BV is not well understood. We tested the hypothesis that having a female partner with BV increases BV-associated bacteria in uncircumcised men. We characterized penile microbiota composition and density (i.e., the quantity of bacteria per swab) by broad-coverage 16S rRNA gene-based sequencing and quantitative PCR (qPCR) in 165 uncircumcised men from Rakai, Uganda. Associations between penile community state types (CSTs) and female partner’s Nugent score were assessed. We found seven distinct penile CSTs of increasing density (CST1 to 7). CST1 to 3 and CST4 to 7 were the two major CST groups. CST4 to 7 had higher prevalence and abundance of BV-associated bacteria, such as <em>Mobiluncus</em> and <em>Dialister</em>, than CST1 to 3. Men with CST4 to 7 were significantly more likely to have a female partner with a high Nugent score (<em>P</em> = 0.03). Men with two or more extramarital partners were significantly more likely to have CST4 to 7 than men with only marital partners (CST4 to 7 prevalence ratio, 1.84; 95% confidence interval [CI], 1.16 to 2.92). Female partner Nugent BV is significantly associated with penile microbiota. Our data support the exchange of BV-associated bacteria through intercourse, which may explain BV recurrence and persistence.
Liu CM, Price LB, Hungate BA, Abraham AG, Larsen LA, Christensen K, Stegger M, Skov R, and Andersen PS (2015) Staphylococcus aureus and the ecology of the nasal microbiome. Science Advances 1(5): e1400216.
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Read PublicationThe human microbiome can play a key role in host susceptibility to pathogens, including in the nasal cavity, a site favored by <em>Staphylococcus aureus</em>. However, what determines our resident nasal microbiota—the host or the environment—and can interactions among nasal bacteria determine <em>S. aureus</em> colonization? Our study of 46 monozygotic and 43 dizygotic twin pairs revealed that nasal microbiota is an environmentally derived trait, but the host’s sex and genetics significantly influence nasal bacterial density. Although specific taxa, including lactic acid bacteria, can determine <em>S. aureus</em> colonization, their negative interactions depend on thresholds of absolute abundance. These findings demonstrate that nasal microbiota is not fixed by host genetics and opens the possibility that nasal microbiota may be manipulated to prevent or eliminate <em>S. aureus</em> colonization.
Liu X-JA, Fike JH, Galbraith JM, Fike WB (2015) Biosolids Amendment and Harvest Frequency Affect Nitrogen Use Dynamics of Switchgrass Grown for Biofuel Production. BioEnergy Research 8(2): 560-569.
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Read PublicationNitrogen use efficiency (NUE) is a crucial index for developing sustainable bioenergy cropping systems. The objective of this study was to examine switchgrass (<em class="EmphasisTypeItalic ">Panicum virgatum</em> L.) NUE by using a low-cost organic amendment under different harvest frequencies. Aerobically digested biosolids were applied at 0, 153, 306, and 459 kg N ha<sup>−1</sup> in a small plot study, and lime-stabilized biosolids were applied at 0, 77, and 154 kg N ha<sup>−1</sup> in a field-scale study in Virginia, USA. Switchgrass was harvested once or twice per season. Switchgrass N concentration and N removal were measured to estimate switchgrass NUE, annual N recovery (ANR), and partial factor productivity (PFP). Across N rates, biosolid application increased biomass N concentration and removal by 29 % and 84 % and decreased NUE, ANR, and PFP in the plot study, but effects were inconsistent in the field study. Low NUE, ANR, and PFP obtained with a single, end-of-season harvest were likely functions of low feedstock N concentrations. Switchgrass harvested in summer had highest N concentrations. Cutting twice per season removed more N than cutting once; the resulting increase in NUE reflects differences in feedstock N concentrations rather than differences in yield. Our results suggest that biosolids can be applied as an alternative N source to support plant growth, and cutting once per season is preferable in sustainable biofuel production systems.
Liu X-JA, Fike JH, Galbraith JM, Fike WB, Parrish DJ, Evanylo GK, Strahm BD (2015) Effects of harvest frequency and biosolids application on switchgrass yield, feedstock quality, and theoretical ethanol yield. GCB Bioenergy 7(1): 112-121.
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Read PublicationSustainable development of a bioenergy industry will require low-cost, high-yielding biomass feedstock of desirable quality. Switchgrass (<em>Panicum virgatum</em> L.) is one of the primary feedstock candidates in North America, but the potential to grow this biomass crop using fertility from biosolids has not been fully explored. The objective of this study was to examine the effects of harvest frequency and biosolids application on switchgrass in Virginia, USA. ‘Cave-in-Rock’ switchgrass from well-established plots was cut once (November) or twice (July and November) per year between 2010 and 2012. Class A biosolids were applied once at rates of 0, 153, 306, and 459 kg N ha<sup>−1</sup> in May 2010. Biomass yield, neutral and acid detergent fiber, cellulose, hemicellulose, lignin, and ash were determined. Theoretical ethanol potential (TEP, l ethanol Mg<sup>−1</sup> biomass) and yield (TEY, l ethanol ha<sup>−1</sup>) were calculated based on cellulose and hemicellulose concentrations. Cutting twice per season produced greater biomass yields than one cutting (11.7 vs. 9.8 Mg ha<sup>−1</sup>) in 2011, but no differences were observed in other years. Cutting once produced feedstock with greater TEP (478 vs. 438 l Mg<sup>−1</sup>), but no differences in TEY between cutting frequencies. Biosolids applied at 153, 306, and 459 kg N ha<sup>−1</sup> increased biomass yields by 25%, 37%, and 46%, and TEY by 25%, 34%, and 42%, respectively. Biosolids had inconsistent effects on feedstock quality and TEP. A single, end-of-season harvest likely will be preferred based on apparent advantages in feedstock quality. Biosolids can serve as an effective alternative to N fertilizer in switchgrass-to-energy systems.
Lundy ME, Pittelkow CM, Linquist BA, Liang X, van Groenigen KJ, Lee J, Six J, Venterea RT, van Kessel C (2015) Nitrogen fertilization reduces yield declines following no-till adoption. Field Crops Research 183: 204-210.
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Read PublicationConservation agriculture (CA) has been promoted as a method of sustainable intensification and climate change mitigation and is being widely practiced and implemented globally. However, no-till (NT), a fundamental component of CA, has been shown to reduce yields in many cases. In order to maintain yields following adoption of CA, it has been recently suggested that fertilizer application should be an integral component of CA. To determine the contribution of nitrogen (N) fertilizer in minimizing yield declines following NT implementation, we assessed 2759 paired comparisons of NT and conventional tillage (CT) systems from 325 studies reported in the peer-reviewed literature between 1980 and 2013. Overall, we found that NT yields decreased −10.7% (−14.8% to −6.5%) and −3.7% (−5.3% to −2.2%) relative to CT in tropical/subtropical and temperate regions, respectively. Among management and environmental variables that included: the rate of N fertilization; the duration of the NT/CT comparison; residue, rotation, and irrigation practices; the crop type; and the site aridity, N rate was the most important explanatory variable for NT yield declines in tropical/subtropical regions. In temperate regions, N fertilization rates were relatively less important. NT yield declines were most consistently observed at low rates of N fertilization during the first 2 years of NT adoption in tropical/subtropical regions. Applications of N fertilizer at rates of up to 85 ± 12 kg N ha<sup>−1</sup> yr<sup>−1</sup> significantly reduced NT yield declines in these scenarios. While this result should not be viewed as a rate recommendation, it does suggest that farmers applying rates of N fertilizer that are low for their specific system will, on average, see higher NT yields if they increase application rates. In addition, when crop rotation was not practiced or residues were removed from the field, NT yield declines were magnified by low rates of N fertilization in tropical/subtropical regions. These results, based on a global data set and across a broad range of crops, highlight the importance of N fertilization in counteracting yield declines in NT systems, particularly in tropical/subtropical regions.
Martin KL, Hurteau MD, Hungate BA, Koch GW, North MP (2015) Carbon Tradeoffs of Restoration and Provision of Endangered Species Habitat in a Fire-Maintained Forest. Ecosystems 18(1): 76-88.
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Read PublicationForests are a significant part of the global carbon cycle and are increasingly viewed as tools for mitigating climate change. Natural disturbances, such as fire, can reduce carbon storage. However, many forests and dependent species evolved with frequent fire as an integral ecosystem process. We used a landscape forest simulation model to evaluate the effects of endangered species habitat management on carbon sequestration. We compared unmanaged forests (control) to forests managed with prescribed burning and prescribed burning combined with thinning. Management treatments followed guidelines of the recovery plan for the endangered red-cockaded woodpecker (RCW), which requires low-density longleaf pine (Pinus palustris) forest. The unmanaged treatment provided the greatest carbon storage, but at the cost of lost RCW habitat. Thinning and burning treatments expanded RCW habitat by increasing the dominance of longleaf pine and reducing forest density, but stored 22% less total ecosystem carbon compared to the control. Our results demonstrate that continued carbon sequestration and the provision of RCW habitat are not incompatible goals, although there is a tradeoff between habitat extent and total ecosystem carbon across the landscape. Management for RCW habitat might also increase ecosystem resilience, as longleaf pine is tolerant of fire and drought, and resistant to pests. Restoring fire-adapted forests requires a reduction in carbon. However, the size of the reduction, the effects on sequestration rates, and the co-benefits from other ecosystem services should be evaluated in the context of the specific forest community targeted for restoration
Mau RL, Liu CM, Aziz M, Schwartz E, Dijkstra P, Marks JC, Price LB, Keim P, and Hungate BA (2015) Linking soil bacterial biodiversity and soil carbon stability. The ISME Journal 9(6): 1477–1480.
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Read PublicationNative soil carbon (C) can be lost in response to fresh C inputs, a phenomenon observed for decades yet still not understood. Using dual-stable isotope probing, we show that changes in the diversity and composition of two functional bacterial groups occur with this ‘priming’ effect. A single-substrate pulse suppressed native soil C loss and reduced bacterial diversity, whereas repeated substrate pulses stimulated native soil C loss and increased diversity. Increased diversity after repeated C amendments contrasts with resource competition theory, and may be explained by increased predation as evidenced by a decrease in bacterial 16S rRNA gene copies. Our results suggest that biodiversity and composition of the soil microbial community change in concert with its functioning, with consequences for native soil C stability.
Mayor JR, Mack MC, Schuur EAG (2015) Decoupled stoichiometric, isotopic, and fungal responses of an ectomycorrhizal black spruce forest to nitrogen and phosphorus additions. Soil Biology and Biochemistry 88: 247–256.
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Read PublicationMany northern forests are limited by nitrogen (N) availability, slight changes in which can have profound effects on ecosystem function and the activity of ectomycorrhizal (EcM) fungi. Increasing N and phosphorus (P) availability, an analog to accelerated soil organic matter decomposition in a warming climate, could decrease plant dependency on EcM fungi and increase plant productivity as a result of greater carbon use efficiency. However, the impact of altered N and P availability on the growth and activity of EcM fungi in boreal forests remains poorly understood despite recognition of their importance to host plant nutrition and soil carbon sequestration. To address such uncertainty we examined above and belowground ecosystem properties in a boreal black spruce forest following five years of factorial N and P additions. By combining detailed soil, fungal, and plant <em>δ</em><sup>15</sup>N measurements with <em>in situ</em> metrics of fungal biomass, growth, and activity, we found both expected and unexpected patterns. Soil nitrate isotope values became <sup>15</sup>N enriched in response to both N and P additions; fungal biomass was repressed by N yet both biomass and growth were stimulated by P; and, black spruce dependency on EcM derived N increased slightly when N and P were added alone yet significantly declined when added in combination. These findings contradict predictions that N fertilization would increase plant P demands and P fertilization would further exacerbate plant N demands. As a result, the prediction that EcM fungi predictably respond to plant N limitation was not supported. These findings highlight P as an under appreciated mediator of the activity of denitrifying bacteria, EcM fungi, and the dynamics of N cycles in boreal forests. Further, use of <em>δ</em><sup>15</sup>N values from bulk soils, plants, and fungi to understand how EcM systems respond to changing nutrient availabilities will often require additional ecological information.
McHugh TA, Morrissey EM, Reed SC, Hungate BA, Schwartz E (2015) Water from air: an overlooked source of moisture in arid and semiarid regions. Scientific Reports 5: 13767.
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Read PublicationWater drives the functioning of Earth’s arid and semiarid lands. Drylands can obtain water from sources other than precipitation, yet little is known about how non-rainfall water inputs influence dryland communities and their activity. In particular, water vapor adsorption – movement of atmospheric water vapor into soil when soil air is drier than the overlying air – likely occurs often in drylands, yet its effects on ecosystem processes are not known. By adding <sup>18</sup>O-enriched water vapor to the atmosphere of a closed system, we documented the conversion of water vapor to soil liquid water across a temperature range typical of arid ecosystems. This phenomenon rapidly increased soil moisture and stimulated microbial carbon (C) cycling, and the flux of water vapor to soil had a stronger impact than temperature on microbial activity. In a semiarid grassland, we also observed that non-rainfall water inputs stimulated microbial activity and C cycling. Together these data suggest that, during rain-free periods, atmospheric moisture in drylands may significantly contribute to variation in soil water content, thereby influencing ecosystem processes. The simple physical process of adsorption of water vapor to soil particles, forming liquid water, represents an overlooked but potentially important contributor to C cycling in drylands.
Melvin AM, Mack MC, Johnstone JF, McGuire AD, Genet H, Schuur EAG (2015) Differences in Ecosystem Carbon Distribution and Nutrient Cycling Linked to Forest Tree Species Composition in a Mid-Successional Boreal Forest. Ecosystems 18 (8): 1472-1488.
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Read PublicationIn the boreal forest of Alaska, increased fire severity associated with climate change is expanding deciduous forest cover in areas previously dominated by black spruce (<em class="EmphasisTypeItalic ">Picea mariana</em>). Needle-leaf conifer and broad-leaf deciduous species are commonly associated with differences in tree growth, carbon (C) and nutrient cycling, and C accumulation in soils. Although this suggests that changes in tree species composition in Alaska could impact C and nutrient pools and fluxes, few studies have measured these linkages. We quantified C, nitrogen, phosphorus, and base cation pools and fluxes in three stands of black spruce and Alaska paper birch (<em class="EmphasisTypeItalic ">Betula neoalaskana</em>) that established following a single fire event in 1958. Paper birch consistently displayed characteristics of more rapid C and nutrient cycling, including greater aboveground net primary productivity, higher live foliage and litter nutrient concentrations, and larger ammonium and nitrate pools in the soil organic layer (SOL). Ecosystem C stocks (aboveground + SOL + 0–10 cm mineral soil) were similar for the two species; however, in black spruce, 78% of measured C was found in soil pools, primarily in the SOL, whereas aboveground biomass dominated ecosystem C pools in birch forest. Radiocarbon analysis indicated that approximately one-quarter of the black spruce SOL C accumulated prior to the 1958 fire, whereas no pre-fire C was observed in birch soils. Our findings suggest that tree species exert a strong influence over C and nutrient cycling in boreal forest and forest compositional shifts may have long-term implications for ecosystem C and nutrient dynamics.
Morrissey EM, Mchugh TA, Preteska L, Hayer M, Dijkstra P, Hungate BA, Schwartz E (2015) Dynamics of extracellular DNA decomposition and bacterial community composition in soil. Soil Biology and Biochemistry 86: 42-49.
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Read PublicationMicrobial necromass is an important source of stabilized organic matter in soil, yet the decomposition dynamics of necromass constituents have not been adequately characterized. This includes DNA, a nutrient-rich molecule that when released into the environment as extracellular DNA (eDNA) can be readily used by soil microorganisms. However, the ecological relevance of eDNA as a nutrient source for soil microorganisms is relatively unknown. To address these deficits, we performed a laboratory experiment wherein soils were amended with <sup>13</sup>C-labeled eDNA and clay minerals known to interact with DNA (kaolinite and montmorillonite). The amount of eDNA-carbon remaining in the soil declined exponentially over time. Kaolinite amendment decreased eDNA decomposition rates and, after 30 days, retained a higher fraction of eDNA-carbon (∼70% remaining) than control or montmorillonite soils (∼40% remaining), indicating that clay mineral sorption can stabilize eDNA-derived carbon in soil. Sequencing of bacterial 16S rRNA genes showed that during the incubation the relative abundance of the added eDNA's sequence decreased by 98%, 92% and 99% in the control, montmorillonite, and kaolinite amended soils respectively. These results suggest that the fraction of eDNA-carbon that remained in the soil was incorporated into microbial biomass, firmly bound to soil constituents, or fragmented and no longer amenable to sequencing. In addition, the eDNA amendment affected the composition of the bacterial community. Specifically, the relative abundance of select phyla (Planctomycetes and TM7) and genera (e.g., <em>Arthrobacter</em> and <em>Nocardioides</em>) were elevated in soils that received eDNA, suggesting these groups may be particularly effective at degrading eDNA and using it for growth. Taken together, these results indicate that while eDNA is consumed by bacteria in soil, a fraction of eDNA material is resistant to decomposition, particularly when stabilized by soil minerals, suggesting a substantial amount of recalcitrant eDNA could accumulate over time.
Natali SM, Schuur EaG, Mauritz M, Schade JD, Celis G, Crummer KG, Johnston C, Krapek J, Pegoraro E, Salmon VG, Webb EE (2015) Permafrost thaw and soil moisture driving CO2 and CH4 release from upland tundra. Journal of Geophysical Research: Biogeosciences 120(3): 525-537.
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Read PublicationAs permafrost degrades, the amount of organic soil carbon (C) that thaws during the growing season will increase, but decomposition may be limited by saturated soil conditions common in high-latitude ecosystems. However, in some areas, soil drying is expected to accompany permafrost thaw as a result of increased water drainage, which may enhance C release to the atmosphere. We examined the effects of ecosystem warming, permafrost thaw, and soil moisture changes on C balance in an upland tundra ecosystem. This study was conducted at a water table drawdown experiment, established in 2011 and located within the Carbon in Permafrost Experimental Heating Research project, an ecosystem warming and permafrost thawing experiment in Alaska. Warming and drying increased cumulative growing season ecosystem respiration by ~20% over 3 years of this experiment. Warming caused an almost twofold increase in decomposition of a common substrate in surface soil (0–10 cm) across all years, and drying caused a twofold increase in decomposition (0–20 cm) relative to control after 3 years of drying. Decomposition of older C increased in the dried and in the combined warmed + dried plots based on soil pore space <sup>14</sup>CO<sub>2</sub>. Although upland tundra systems have been considered CH<sub>4</sub> sinks, warming and ground thaw significantly increased CH<sub>4</sub> emission rates. Water table depth was positively correlated with monthly respiration and negatively correlated with CH<sub>4</sub> emission rates. These results demonstrate that warming and drying may increase loss of old permafrost C from tundra ecosystems, but the form and magnitude of C released to the atmosphere will be driven by changes in soil moisture.
Ogle K, Barber JJ, Barron-Gafford GA, Bentley LP, Cable JM, Huxman TE, Loik ME, Tissue DT (2015) Quantifying ecological memory in plant and ecosystem processes. Ecology Letters 18(3): 221-235.
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Read PublicationThe role of time in ecology has a long history of investigation, but ecologists have largely restricted their attention to the influence of concurrent abiotic conditions on rates and magnitudes of important ecological processes. Recently, however, ecologists have improved their understanding of ecological processes by explicitly considering the effects of antecedent conditions. To broadly help in studying the role of time, we evaluate the length, temporal pattern, and strength of memory with respect to the influence of antecedent conditions on current ecological dynamics. We developed the stochastic antecedent modelling (SAM) framework as a flexible analytic approach for evaluating exogenous and endogenous process components of memory in a system of interest. We designed SAM to be useful in revealing novel insights promoting further study, illustrated in four examples with different degrees of complexity and varying time scales: stomatal conductance, soil respiration, ecosystem productivity, and tree growth. Models with antecedent effects explained an additional 18–28% of response variation compared to models without antecedent effects. Moreover, SAM also enabled identification of potential mechanisms that underlie components of memory, thus revealing temporal properties that are not apparent from traditional treatments of ecological time-series data and facilitating new hypothesis generation and additional research.
Ogle K., Pendall E (2015) Isotope partitioning of soil respiration: A Bayesian solution to accommodate multiple sources of variability. Journal of Geophysical Research: Biogeosciences 120(2): 221-236.
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Read Publication<section id="abstract" class="article-section article-section--abstract article-tools__article-section--is-active">
<div class="article-section__content mainAbstract">
Isotopic methods offer great potential for partitioning trace gas fluxes such as soil respiration into their different source contributions. Traditional partitioning methods face challenges due to variability introduced by different measurement methods, fractionation effects, and end-member uncertainty. To address these challenges, we describe a hierarchical Bayesian (HB) approach for isotopic partitioning of soil respiration that directly accommodates such variability. We apply our HB method to data from an experiment conducted in a shortgrass steppe ecosystem, where decomposition was previously shown to be stimulated by elevated CO<sub>2</sub>. Our approach simultaneously fits Keeling plot (KP) models to observations of soil or soil-respired <em>δ</em><sup>13</sup>C and [CO<sub>2</sub>] obtained via chambers and gas wells, corrects the KP intercepts for apparent fractionation (Δ) due to isotope-specific diffusion rates and/or method artifacts, estimates method- and treatment-specific values for Δ, propagates end-member uncertainty, and calculates proportional contributions from two distinct respiration sources (“old” and “new” carbon). The chamber KP intercepts were estimated with greater confidence than the well intercepts and compared to the theoretical value of 4.4‰, our results suggest that Δ varies between 2 and 5.2‰ depending on method (chambers versus wells) and CO<sub>2</sub> treatment. Because elevated CO<sub>2</sub> plots were fumigated with <sup>13</sup>C-depleted CO<sub>2</sub>, the source contributions were tightly constrained, and new C accounted for 64% (range = 55–73%) of soil respiration. The contributions were less constrained for the ambient CO<sub>2</sub> treatments, but new C accounted for significantly less (47%, range = 15–82%) of soil respiration. Our new HB partitioning approach contrasts our original analysis (higher contribution of old C under elevated CO<sub>2</sub>) because it uses additional data sources, accounts for end-member bias, and estimates apparent fractionation effects.
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Penton CR, St Louis D, Pham A, Cole JR, Wu L, Luo Y, Schuur EAG, Zhou J, Tiedje JM (2015) Denitrifying and diazotrophic community responses to artificial warming in permafrost and tallgrass prairie soils. Frontiers in Microbiology 6: 746.
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Read PublicationIncreasing temperatures have been shown to impact soil biogeochemical processes, although the corresponding changes to the underlying microbial functional communities are not well understood. Alterations in the nitrogen (N) cycling functional component are particularly important as N availability can affect microbial decomposition rates of soil organic matter and influence plant productivity. To assess changes in the microbial component responsible for these changes, the composition of the N-fixing (<i>nifH</i>), and denitrifying (<i>nirS, nirK, nosZ</i>) soil microbial communities was assessed by targeted pyrosequencing of functional genes involved in N cycling in two major biomes where the experimental effect of climate warming is under investigation, a tallgrass prairie in Oklahoma (OK) and the active layer above permafrost in Alaska (AK). Raw reads were processed for quality, translated with frameshift correction, and a total of 313,842 amino acid sequences were clustered and linked to a nearest neighbor using reference datasets. The number of OTUs recovered ranged from 231 (NifH) to 862 (NirK). The N functional microbial communities of the prairie, which had experienced a decade of experimental warming were the most affected with changes in the richness and/or overall structure of NifH, NirS, NirK and NosZ. In contrast, the AK permafrost communities, which had experienced only 1 year of warming, showed decreased richness and a structural change only with the <i>nirK</i>-harboring bacterial community. A highly divergent <i>nirK</i>-harboring bacterial community was identified in the permafrost soils, suggesting much novelty, while other N functional communities exhibited similar relatedness to the reference databases, regardless of site. Prairie and permafrost soils also harbored highly divergent communities due mostly to differing major populations.
Pittelkow CM, Liang X, Linquist BA, Van Groenigen KJ, Lee J, Lundy ME, Van Gestel N, Six J, Venterea RT, Van Kessel C (2015) Productivity limits and potentials of the principles of conservation agriculture. Nature 517(7534): 365-368.
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Read PublicationOne of the primary challenges of our time is to feed a growing and more demanding world population with reduced external inputs and minimal environmental impacts, all under more variable and extreme climate conditions in the future<sup><a id="ref-link-1" title="Foley, J. A. et al. Solutions for a cultivated planet. Nature 478, 337-342 (2011)" href="http://www.nature.com/nature/journal/v517/n7534/full/nature13809.html#ref1">1</a>, <a id="ref-link-2" title="Lobell, D. B. et al. Prioritizing climate change adaptation needs for food security in 2030. Science 319, 607-610 (2008)" href="http://www.nature.com/nature/journal/v517/n7534/full/nature13809.html#ref2">2</a>, <a id="ref-link-3" title="Godfray, H. C. J. & Garnett, T. Food security and sustainable intensification. Phil. Trans. R. Soc. B 369, 20120273 (2014)" href="http://www.nature.com/nature/journal/v517/n7534/full/nature13809.html#ref3">3</a>, <a id="ref-link-4" title="Tilman, D., Balzer, C., Hill, J. & Befort, B. L. Global food demand and the sustainable intensification of agriculture. Proc. Natl Acad. Sci. USA 108, 20260-20264 (2011)" href="http://www.nature.com/nature/journal/v517/n7534/full/nature13809.html#ref4">4</a></sup>. Conservation agriculture represents a set of three crop management principles that has received strong international support to help address this challenge<sup><a id="ref-link-5" title="FAO. Save and Grow: A Policymaker/'s Guide to the Sustainable Intensification of Smallholder Crop Production 1-37 (FAO, 2011)" href="http://www.nature.com/nature/journal/v517/n7534/full/nature13809.html#ref5">5</a>, <a id="ref-link-6" title="Hobbs, P. R., Sayre, K. & Gupta, R. The role of conservation agriculture in sustainable agriculture. Phil. Trans. R. Soc. B 363, 543-555 (2008)" href="http://www.nature.com/nature/journal/v517/n7534/full/nature13809.html#ref6">6</a></sup>, with recent conservation agriculture efforts focusing on smallholder farming systems in sub-Saharan Africa and South Asia<sup><a id="ref-link-7" title="Stevenson, J. R., Serraj, R. & Cassman, K. G. Evaluating conservation agriculture for small-scale farmers in sub-Saharan Africa and South Asia. Agric. Ecosyst. Environ. 187, 1-10 (2014)" href="http://www.nature.com/nature/journal/v517/n7534/full/nature13809.html#ref7">7</a></sup>. However, conservation agriculture is highly debated, with respect to both its effects on crop yields<sup><a id="ref-link-8" title="Giller, K. E., Witter, E., Corbeels, M. & Tittonell, P. Conservation agriculture and smallholder farming in Africa: the heretics/' view. Field Crops Res. 114, 23-34 (2009)" href="http://www.nature.com/nature/journal/v517/n7534/full/nature13809.html#ref8">8</a>, <a id="ref-link-9" title="Rusinamhodzi, L. et al. A meta-analysis of long-term effects of conservation agriculture on maize grain yield under rain-fed conditions. Agron. Sust. Dev. 31, 657-673 (2011)" href="http://www.nature.com/nature/journal/v517/n7534/full/nature13809.html#ref9">9</a>, <a id="ref-link-10" title="Brouder, S. M. & Gomez-Macpherson, H. The impact of conservation agriculture on smallholder agricultural yields: a scoping review of the evidence. Agric. Ecosyst. Environ. 187, 11-32 (2014)" href="http://www.nature.com/nature/journal/v517/n7534/full/nature13809.html#ref10">10</a></sup> and its applicability in different farming contexts<sup><a id="ref-link-11" title="Stevenson, J. R., Serraj, R. & Cassman, K. G. Evaluating conservation agriculture for small-scale farmers in sub-Saharan Africa and South Asia. Agric. Ecosyst. Environ. 187, 1-10 (2014)" href="http://www.nature.com/nature/journal/v517/n7534/full/nature13809.html#ref7">7</a>, <a id="ref-link-12" title="Andersson, J. A. & Giller, K. E. in Contested Agronomy: Agricultural Research in a Changing World (eds Sumberg, J. & Thompson, J.) Ch. 2 22-46 (Earthscan, 2012)" href="http://www.nature.com/nature/journal/v517/n7534/full/nature13809.html#ref11">11</a>, <a id="ref-link-13" title="Giller, K. E. et al. A research agenda to explore the role of conservation agriculture in African smallholder farming systems. Field Crops Res. 124, 468-472 (2011)" href="http://www.nature.com/nature/journal/v517/n7534/full/nature13809.html#ref12">12</a>, <a id="ref-link-14" title="Friedrich, T., Derpsch, R. & Kassam, A. Overview of the global spread of conservation agriculture. Field Actions Sci. Rep. 6, 1941 (2012)" href="http://www.nature.com/nature/journal/v517/n7534/full/nature13809.html#ref13">13</a></sup>. Here we conduct a global meta-analysis using 5,463 paired yield observations from 610 studies to compare no-till, the original and central concept of conservation agriculture, with conventional tillage practices across 48 crops and 63 countries. Overall, our results show that no-till reduces yields, yet this response is variable and under certain conditions no-till can produce equivalent or greater yields than conventional tillage. Importantly, when no-till is combined with the other two conservation agriculture principles of residue retention and crop rotation, its negative impacts are minimized. Moreover, no-till in combination with the other two principles significantly increases rainfed crop productivity in dry climates, suggesting that it may become an important climate-change adaptation strategy for ever-drier regions of the world. However, any expansion of conservation agriculture should be done with caution in these areas, as implementation of the other two principles is often challenging in resource-poor and vulnerable smallholder farming systems, thereby increasing the likelihood of yield losses rather than gains. Although farming systems are multifunctional, and environmental and socio-economic factors need to be considered<sup><a id="ref-link-15" title="Godfray, H. C. et al. Food security: the challenge of feeding 9 billion people. Science 327, 812-818 (2010)" href="http://www.nature.com/nature/journal/v517/n7534/full/nature13809.html#ref14">14</a>, <a id="ref-link-16" title="Sachs, J. et al. Monitoring the world/'s agriculture. Nature 466, 558-560 (2010)" href="http://www.nature.com/nature/journal/v517/n7534/full/nature13809.html#ref15">15</a>, <a id="ref-link-17" title="Palm, C., Blanco-Canqui, H., DeClerck, F., Gatere, L. & Grace, P. Conservation agriculture and ecosystem services: An overview. Agric. Ecosyst. Environ. 187, 87-105 (2014)" href="http://www.nature.com/nature/journal/v517/n7534/full/nature13809.html#ref16">16</a></sup>, our analysis indicates that the potential contribution of no-till to the sustainable intensification of agriculture is more limited than often assumed.
Pittelkow CM, Linquist BA, Lundy ME, Liang X, van Groenigen KJ, Lee J, van Gestel N, Six J, Venterea RT, van Kessel C (2015) When does no-till yield more? A global meta-analysis. Field Crops Research 183: 156–168.
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Read PublicationNo-till agriculture represents a relatively widely adopted management system that aims to reduce soil erosion, decrease input costs, and sustain long-term crop productivity. However, its impacts on crop yields are variable, and an improved understanding of the factors limiting productivity is needed to support evidence-based management decisions. We conducted a global meta-analysis to evaluate the influence of various crop and environmental variables on no-till relative to conventional tillage yields using data obtained from peer-reviewed publications (678 studies with 6005 paired observations, representing 50 crops and 63 countries). Side-by-side yield comparisons were restricted to studies comparing conventional tillage to no-till practices in the absence of other cropping system modifications. Crop category was the most important factor influencing the overall yield response to no-till followed by aridity index, residue management, no-till duration, and N rate. No-till yields matched conventional tillage yields for oilseed, cotton, and legume crop categories. Among cereals, the negative impacts of no-till were smallest for wheat (−2.6%) and largest for rice (−7.5%) and maize (−7.6%). No-till performed best under rainfed conditions in dry climates, with yields often being equal to or higher than conventional tillage practices. Yields in the first 1–2 years following no-till implementation declined for all crops except oilseeds and cotton, but matched conventional tillage yields after 3–10 years except for maize and wheat in humid climates. Overall, no-till yields were reduced by 12% without N fertilizer addition and 4% with inorganic N addition. Our study highlights factors contributing to and/or decreasing no-till yield gaps and suggests that improved targeting and adaptation, possibly including additional system modifications, are necessary to optimize no-till performance and contribute to food production goals. In addition, our results provide a basis for conducting trade-off analyses to support the development of no-till crop management and international development strategies based on available scientific evidence.
Price LB, Koch BJ, Hungate BA (2015) Ominous projections for global antibiotic use in food-animal production. Proceedings of the National Academy of Sciences 112 (18): 5554-5555.
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Read Publication<p id="p-2" class="flushleft">Alexander Fleming famously warned that the ignorant may someday misuse his life-saving discovery—penicillin—and select for resistant bacteria (<a id="xref-ref-1-1" class="xref-bibr" href="http://www.pnas.org/content/112/18/5554.full#ref-1">1</a>). This was prescient given the widespread use of subtherapeutic antibiotics by food-animal producers today. According to the findings of Van Boeckel et al. (<a id="xref-ref-2-1" class="xref-bibr" href="http://www.pnas.org/content/112/18/5554.full#ref-2">2</a>) in PNAS, the proliferation of ignorance is only poised to increase. Using global datasets of veterinary antibiotic use, livestock densities, and economic projections of meat demand, Van Boeckel et al. (<a id="xref-ref-2-2" class="xref-bibr" href="http://www.pnas.org/content/112/18/5554.full#ref-2">2</a>) estimate that from 2010 to 2030 antibiotic use in food-animal production will increase by 67%, from 63,151 ± 1,560 tons to 105,596 ± 3,605 tons.</p>
<p id="p-3">The study by Van Boeckel et al. (<a id="xref-ref-2-3" class="xref-bibr" href="http://www.pnas.org/content/112/18/5554.full#ref-2">2</a>) is the first to estimate global use of antibiotics in livestock production, and to disaggregate that global figure into estimates for each of 228 countries. However, their estimate is based on data from only 32 countries. Using a clear framework and a state-of-the-art Bayesian statistical model, the authors extrapolate from the most reliable data available to arrive at the global sum. This is an admirable approach to a difficult problem, but it raises a question: Why not derive the values more simply, by summing data from all 228 countries, using the actual records of antibiotic use in livestock production? After all, this is how we quantify global fossil fuel use (<a id="xref-ref-3-1" class="xref-bibr" href="http://www.pnas.org/content/112/18/5554.full#ref-3">3</a>), livestock production and trade (<a id="xref-ref-4-1" class="xref-bibr" href="http://www.pnas.org/content/112/18/5554.full#ref-4">4</a>, <a id="xref-ref-5-1" class="xref-bibr" href="http://www.pnas.org/content/112/18/5554.full#ref-5">5</a>), and the use of fertilizers in agriculture (<a id="xref-ref-4-2" class="xref-bibr" href="http://www.pnas.org/content/112/18/5554.full#ref-4">4</a>). For many assessments of global economic activity, including these, the actual data exist. However, for antibiotics in livestock production, a statistical model is the best option because comprehensive data on the use of antibiotics in livestock production are not available. Most countries do not record the sale and use of antibiotics, in part because practitioners may be reluctant to release those data. Despite this limitation, Van Boeckel et al. (<a id="xref-ref-2-4" class="xref-bibr" href="http://www.pnas.org/content/112/18/5554.full#ref-2">2</a>) provide the first global assessment of antibiotic use in livestock production. Their estimate is important: The figure is large and has been notoriously difficult to extract (<a id="xref-ref-6-1" class="xref-bibr" href="http://www.pnas.org/content/112/18/5554.full#ref-6">6</a>), and it sets the stage for understanding the global impacts of profligate use of these powerful drugs.</p>
Ryan E, Ogle K, Zelikova TJ, LeCain DR, Williams DG, Morgan JA, Pendall E (2015) Antecedent moisture and temperature conditions modulate the response of ecosystem respiration to elevated CO2 and warming. Global Change Biology 21(7): 2588-2602.
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Read PublicationTerrestrial plant and soil respiration, or ecosystem respiration (R<sub>eco</sub>), represents a major CO<sub>2</sub> flux in the global carbon cycle. However, there is disagreement in how R<sub>eco</sub> will respond to future global changes, such as elevated atmosphere CO<sub>2</sub> and warming. To address this, we synthesized six years (2007–2012) of R<sub>eco</sub> data from the Prairie Heating And CO<sub>2</sub> Enrichment (PHACE) experiment. We applied a semi-mechanistic temperature–response model to simultaneously evaluate the response of R<sub>eco</sub> to three treatment factors (elevated CO<sub>2</sub>, warming, and soil water manipulation) and their interactions with antecedent soil conditions [e.g., past soil water content (SWC) and temperature (SoilT)] and aboveground factors (e.g., vapor pressure deficit, photosynthetically active radiation, vegetation greenness). The model fits the observed R<sub>eco</sub> well (<em>R</em><sup>2 </sup>= 0.77). We applied the model to estimate annual (March–October) R<sub>eco</sub>, which was stimulated under elevated CO<sub>2</sub> in most years, likely due to the indirect effect of elevated CO<sub>2</sub> on SWC. When aggregated from 2007 to 2012, total six-year R<sub>eco</sub> was stimulated by elevated CO<sub>2</sub> singly (24%) or in combination with warming (28%). Warming had little effect on annual R<sub>eco</sub> under ambient CO<sub>2</sub>, but stimulated it under elevated CO<sub>2</sub> (32% across all years) when precipitation was high (e.g., 44% in 2009, a ‘wet’ year). Treatment-level differences in R<sub>eco</sub> can be partly attributed to the effects of antecedent SoilT and vegetation greenness on the apparent temperature sensitivity of R<sub>eco</sub> and to the effects of antecedent and current SWC and vegetation activity (greenness modulated by VPD) on R<sub>eco</sub> base rates. Thus, this study indicates that the incorporation of both antecedent environmental conditions and aboveground vegetation activity are critical to predicting R<sub>eco</sub> at multiple timescales (subdaily to annual) and under a future climate of elevated CO<sub>2</sub> and warming.
Schuur EAG, McGuire AD, Schädel C, Grosse G, Harden JW, Hayes DJ, Hugelius G, Koven CD, Kuhry P, Lawrence DM, Natali SM, Olefeldt D, Romanovsky VE, Schaefer K, Turetsky MR, Treat CC, Vonk JE (2015) Climate change and the permafrost carbon feedback. Nature 520(7546): 171-179.
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Read PublicationLarge quantities of organic carbon are stored in frozen soils (permafrost) within Arctic and sub-Arctic regions. A warming climate can induce environmental changes that accelerate the microbial breakdown of organic carbon and the release of the greenhouse gases carbon dioxide and methane. This feedback can accelerate climate change, but the magnitude and timing of greenhouse gas emission from these regions and their impact on climate change remain uncertain. Here we find that current evidence suggests a gradual and prolonged release of greenhouse gas emissions in a warming climate and present a research strategy with which to target poorly understood aspects of permafrost carbon dynamics.
Sillett SC, Van Pelt R, Kramer RD, Carroll AL, Koch GW (2015) Biomass and growth potential of Eucalyptus regnans up to 100 m tall. Forest Ecology and Management 348: 78-91.
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Read PublicationPredicting tree biomass and growth increments via allometric equations is routine in forestry, but this approach is problematic in old-growth forests unless equations are derived from trees spanning the full size range. Using intensive measurements of 27 standing <em>Eucalyptus regnans</em> trees 61.1–99.8 m tall and 80–430 years old in Tasmania, Victoria, and New Zealand, we develop allometric equations to predict aboveground attributes, including biomass and annual growth increments, of trees >60 m tall using ground-based measurements. Power functions of diameter underestimate biomass growth increments unless measurements are made above buttressing on the lower trunk. Growth anomalies apparent in several trees suggest that wounded <em>E. regnans</em>expend considerable energy to outgrow decay fungi and prevent structural collapse. Despite declining growth efficiency – defined as biomass growth per unit mass of photosynthetic tissues – with increasing tree size and age, biomass growth increments of <em>E. regnans</em> increase as trees enlarge with age until extrinsic forces cause mortality. The largest living <em>E. regnans</em> has an aboveground biomass of 215 Mg and a growth increment of 0.784 Mg year<sup>−1</sup>, not accounting for mass loss due to decay. An even larger <em>E. regnans</em>tree – killed by fire in 2003 – had an aboveground biomass of ∼270 Mg, an estimated growth increment of ∼1 Mg year<sup>−1</sup>, and was ∼480 years old at the end of its life. Prior to a stand-replacing fire in 2009, Australia’s tallest forest had a maximum aboveground biomass of 1504 Mg ha<sup>−1</sup> and a maximum aboveground carbon mass of 706 Mg ha<sup>−1</sup>.
Simonin M, Le Roux X, Poly F, Lerondelle C, Hungate B, Nunan N, Niboyet A (2015) Coupling Between and Among Ammonia Oxidizers and Nitrite Oxidizers in Grassland Mesocosms Submitted to Elevated CO2 and Nitrogen Supply. Microbial Ecology 70(3): 809-818.
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Read PublicationMany studies have assessed the responses of soil microbial functional groups to increases in atmospheric CO2 or N deposition alone and more rarely in combination. However, the effects of elevated CO2 and N on the (de)coupling between different microbial functional groups (e.g., different groups of nitrifiers) have been barely studied, despite potential consequences for ecosystem functioning. Here, we investigated the short-term combined effects of elevated CO2 and N supply on the abundances of the four main microbial groups involved in soil nitrification: ammonia-oxidizing archaea (AOA), ammonia-oxidizing bacteria (AOB), and nitrite-oxidizing bacteria (belonging to the genera Nitrobacter and Nitrospira) in grassland mesocosms. AOB and AOA abundances responded differently to the treatments: N addition increased AOB abundance, but did not alter AOA abundance. Nitrobacter and Nitrospira abundances also showed contrasted responses to the treatments: N addition increased Nitrobacter abundance, but decreased Nitrospira abundance. Our results support the idea of a niche differentiation between AOB and AOA, and between Nitrobacter and Nitrospira. AOB and Nitrobacter were both promoted at high N and C conditions (and low soil water content for Nitrobacter), while AOA and Nitrospira were favored at low N and C conditions (and high soil water content for Nitrospira). In addition, Nitrobacter abundance was positively correlated to AOB abundance and Nitrospira abundance to AOA abundance. Our results suggest that the couplings between ammonia and nitrite oxidizers are influenced by soil N availability. Multiple environmental changes may thus elicit rapid and contrasted responses between and among the soil ammonia and nitrite oxidizers due to their different ecological requirements.
Sugden AM, A Cooper, C Turney, KA Hughen, BW Brook, HG McDonald, … GW Koch, et al. (2015) Climate killed off the megafauna. Science 349(6248): 598.
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Read Publication<div dir="ltr" data-angle="0" data-font-name="g_font_21_0" data-canvas-width="215.6235865445137">The causes of the Pleistocene extinctions of large numbers of megafaunal species in the Northern Hemisphere remain unclear. A range of evidence points to human hunting, climate change, or a combination of both. Using ancient DNA and detailed paleoclimate data, Cooper et al . report a close relationship between Pleistocene megafau-nal extinction events and rapid warming events at the start</div>
<div dir="ltr" data-angle="0" data-font-name="g_font_21_0" data-canvas-width="204.2591669979572">of interstadial periods. Their analysis strengthens the case for climate change as the key driver of megafaunal extinctions, with human impacts playing a secondary role.</div>
Sullivan BW, Nasto MK, Hart SC, Hungate BA (2015) Proximate controls on semiarid soil greenhouse gas fluxes across 3 million years of soil development. Biogeochemistry 125(3): 375-391.
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Read PublicationSoils are important sources and sinks of three greenhouse gases (GHGs): carbon dioxide (CO<sub>2</sub>), methane (CH<sub>4</sub>), and nitrous oxide (N<sub>2</sub>O). However, it is unknown whether semiarid landscapes are important contributors to global fluxes of these gases, partly because our mechanistic understanding of soil GHG fluxes is largely derived from more humid ecosystems. We designed this study with the objective of identifying the important soil physical and biogeochemical controls on soil GHG fluxes in semiarid soils by observing seasonal changes in soil GHG fluxes across a three million year substrate age gradient in northern Arizona. We also manipulated soil nitrogen (N) and phosphorus availability with 7 years of fertilization and used regression tree analysis to identify drivers of unfertilized and fertilized soil GHG fluxes. Similar to humid ecosystems, soil N<sub>2</sub>O flux was correlated with changes in N and water availability and soil CO<sub>2</sub> efflux was correlated with changes in water availability and temperature. Soil CH<sub>4</sub> uptake was greatest in relatively colder and wetter soils. While fertilization had few direct effects on soil CH<sub>4</sub> flux, soil nitrate was an important predictor of soil CH<sub>4</sub> uptake in unfertilized soils and soil ammonium was an important predictor of soil CH<sub>4</sub> uptake in fertilized soil. Like in humid ecosystems, N gas loss via nitrification or denitrification appears to increase with increases in N and water availability during ecosystem development. Our results suggest that, with some exceptions, the drivers of soil GHG fluxes in semiarid ecosystems are often similar to those observed in more humid ecosystems.
Treat CC, Natali SM, Ernakovich J, Iversen CM, Lupascu M, McGuire AD, Norby RJ, Roy Chowdhury T, Richter A, Šantrůčková H, Schädel C, Schuur EAG, Sloan VL, Turetsky MR, Waldrop MP (2015) A pan-Arctic synthesis of CH4 and CO2 production from anoxic soil incubations. Global Change Biology 21(7): 2787–2803.
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Read PublicationPermafrost thaw can alter the soil environment through changes in soil moisture, frequently resulting in soil saturation, a shift to anaerobic decomposition, and changes in the plant community. These changes, along with thawing of previously frozen organic material, can alter the form and magnitude of greenhouse gas production from permafrost ecosystems. We synthesized existing methane (CH4) and carbon dioxide (CO2) production measurements from anaerobic incubations of boreal and tundra soils from the geographic permafrost region in order to evaluate large-scale controls of anaerobic CO2 and CH4 production and compare the relative importance of landcape-level factors (e.g., vegetation type and landscape position), soil properties (e.g., pH, depth and soil type), and soil environmental conditions (e.g., temperature and relative water table position). We found five-fold higher maximum CH4 production per gram soil carbon from organic soils than mineral soils. Maximum CH4 production from soils in the active layer (ground that thaws and refreezes annually) was nearly four times that of permafrost per gram soil carbon, and CH4 production per gram soil carbon was two times greater from sites without permafrost than sites with permafrost. Maximum CH4 and median anaerobic CO2 production decreased with depth, while CO2:CH4 production increased with depth. Maximum CH4 production was highest in soils with herbaceous vegetation and soils that were either consistently or periodically inundated. This synthesis identifies the need to consider biome, landscape position, and vascular/moss vegetation types when modeling CH4 production in permafrost ecosystems and suggests the need for longer-term anaerobic incubations to fully capture CH4 dynamics. Our results demonstrate that as climate warms in arctic and boreal regions, rates of anaerobic CO2 and CH4 production will increase, not only as a result of increased temperature, but also from shifts in vegetation and increased ground saturation that will accompany permafrost thaw. This article is protected by copyright. All rights reserved
van Groenigen KJ, Xia J, Osenberg CW, Luo Y, Hungate BA (2015) Application of a two-pool model to soil carbon dynamics under elevated CO2. Global Change Biology 21(12): 4293–4297 .
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Read PublicationElevated atmospheric CO<sub>2</sub> concentrations increase plant productivity and affect soil microbial communities, with possible consequences for the turnover rate of soil carbon (C) pools and feedbacks to the atmosphere. In a previous analysis (Van Groenigen <em>et al</em>., 2014), we used experimental data to inform a one-pool model and showed that elevated CO<sub>2</sub> increases the decomposition rate of soil organic C, negating the storage potential of soil. However, a two-pool soil model can potentially explain patterns of soil C dynamics without invoking effects of CO<sub>2</sub> on decomposition rates. To address this issue, we refit our data to a two-pool soil C model. We found that CO<sub>2</sub> enrichment increases decomposition rates of both fast and slow C pools. In addition, elevated CO<sub>2</sub> decreased the carbon use efficiency of soil microbes (CUE), thereby further reducing soil C storage. These findings are consistent with numerous empirical studies and corroborate the results from our previous analysis. To facilitate understanding of C dynamics, we suggest that empirical and theoretical studies incorporate multiple soil C pools with potentially variable decomposition rates.
Vogel JG, He D, Jokela EJ, Hockaday W, Schuur EAG (2015) The effect of fertilization levels and genetic deployment on the isotopic signature, constituents, and chemistry of soil organic carbon in managed loblolly pine (Pinus taeda L.) forests. Forest Ecology and Management 355: 91–100.
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Read PublicationSoil organic carbon (SOC) mass and its constituents, chemistry, and isotopic signatures (Δ<sup>14</sup>C, δ<sup>13</sup>C) were examined for two different loblolly pine (<em>Pinus taeda</em> L.) research installations located in north-central Florida. Both studies were designed as split-plots with the whole plots as different levels of fertilization and herbicide application (cultural intensity), and full-sib families of loblolly pine were the splits. The cultural intensities and the families of loblolly pine were different at each site and so each site was analyzed separately. The plantations were aged 9 or 10 years at the time of soil sampling. At both sites, the overall mass of SOC to a depth of 0–30 cm was unresponsive to the level of family growth or cultural intensity and did not show a trend with aboveground biomass. The SOC pool was further separated into live roots and wood; and density fractionation was used to separate the SOC sample into a light fraction (LF) (<1.7 g cm<sup>−3</sup>) and heavy fraction (HF) with the LF dissected further for charcoal and dead roots. Higher fertilization levels generally depressed fine root (<1 mm) biomass, but whether the effect was significant varied with family and soil horizon. The HF was a relatively small component (<5%) of SOC in these sandy textured soils, but at one of the two sites, the HF was significantly increased with more intensive silviculture and for the faster growing family. The Δ<sup>14</sup>C value of the LF-SOC for one slow growing family under low culture (136 ± 11‰) differed from the faster growing low culture plot, and its relationship to the atmospheric Δ<sup>14</sup>C record suggested that the LF-SOC likely originated prior to stand establishment. The LF chemistry was determined with solid-state <sup>13</sup>C nuclear magnetic resonance (NMR) and cultural intensity did not significantly affect SOC chemistry. However, the family effect was significant for carbohydrates at one site, and for lignin and lipids at the other site. Overall, these results suggest that tree genetics in managed forests can influence SOC chemistry and that the relatively small fractions of SOC can change with management intensity; however, the effect of cultural intensity is minimal for the largest components of SOC and there is no clear relationship between SOC dynamics and aboveground production under the management regimes, and stand ages, examined with these two research installations.
Walker XJ, Mack MC, Johnstone JF (2015) Stable carbon isotope analysis reveals widespread drought stress in boreal black spruce forests. Global Change Biology 21(8): 3102-3113.
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Read PublicationUnprecedented rates of climate warming over the past century have resulted in increased forest stress and mortality worldwide. Decreased tree growth in association with increasing temperatures is generally accepted as a signal of temperature-induced drought stress. However, variations in tree growth alone do not reveal the physiological mechanisms behind recent changes in tree growth. Examining stable carbon isotope composition of tree rings in addition to tree growth can provide a secondary line of evidence for physiological drought stress. In this study, we examined patterns of black spruce growth and carbon isotopic composition in tree rings in response to climate warming and drying in the boreal forest of interior Alaska. We examined trees at three nested scales: landscape, toposequence, and a subsample of trees within the toposequence. At each scale, we studied the potential effects of differences in microclimate and moisture availability by sampling on northern and southern aspects. We found that black spruce radial growth responded negatively to monthly metrics of temperature at all examined scales, and we examined ∆<sup>13</sup>C responses on a subsample of trees as representative of the wider region. The negative ∆<sup>13</sup>C responses to temperature reveal that black spruce trees are experiencing moisture stress on both northern and southern aspects. Contrary to our expectations, ∆<sup>13</sup>C from trees on the northern aspect exhibited the strongest drought signal. Our results highlight the prominence of drought stress in the boreal forest of interior Alaska. We conclude that if temperatures continue to warm, we can expect drought-induced productivity declines across large regions of the boreal forest, even for trees located in cool and moist landscape positions.
Wiechmann ML, Hurteau MD, North MP, Koch GW, Jerabkova L (2015) The carbon balance of reducing wildfire risk and restoring process: an analysis of 10-year post-treatment carbon dynamics in a mixed-conifer forest. Climatic Change 132(4): 709-719.
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Read PublicationForests sequester carbon from the atmosphere, helping mitigate climate change. In fire-prone forests, burn events result in direct and indirect emissions of carbon. High fire-induced tree mortality can cause a transition from a carbon sink to source, but thinning and prescribed burning can reduce fire severity and carbon loss when wildfire occurs. However, treatment implementation requires carbon removal and emissions to reduce high-severity fire risk. The carbon removed and emitted during treatment may be resequestered by subsequent tree growth, although there is much uncertainty regarding the length of time required. To assess the long-term carbon dynamics of thinning and burning treatments, we quantified the 10-year post-treatment carbon stocks and 10-year net biome productivity (NBP) from a full-factorial experiment involving three levels of thinning and two levels of burning in a mixed-conifer forest in California’s Sierra Nevada. Our results indicate that (1) the understory thin treatment, that retained large trees, quickly recovered the initial carbon emissions (NBP = 31.4 ± 4.2 Mg C ha<sup>−1</sup>), (2) the carbon emitted from prescribed fire in the burn-only treatment was resequestered within the historical fire return interval (NBP = 32.8 ± 3.5 Mg C ha<sup>−1</sup>), and (3) the most effective treatment for reducing fire risk, understory thin and burn, had negative NBP (−6.0 ± 4.5 Mg C ha<sup>−1</sup>) because of post-fire large tree mortality. Understory thinning and prescribed burning can help stabilize forest carbon and restore ecosystem resilience, but this requires additional emissions beyond only thinning or only burning. Retaining additional mid-sized trees may reduce the carbon impacts of understory thinning and burning.
Wymore AS, Compson ZG, McDowell WH, Potter JD, Hungate BA, Whitham TG, Marks JC, (2015) Leaf-litter leachate is distinct in optical properties and bioavailability to stream heterotrophs. Freshwater Science 34(3): 857-866.
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Read PublicationDissolved organic C (DOC) leached from leaf litter contributes to the C pool of stream ecosystems and affects C cycling in streams. We studied how differences in leaf-litter chemistry affect the optical properties and decomposition of DOC. We used 2 species of cottonwoods (Populus) and their naturally occurring hybrids that differ in leaf-litter phytochemistry and decomposition rate. We measured DOC and nutrient concentration in leaf leachates and determined the effect of DOC quality on heterotrophic respiration in 24-h incubations with stream sediments. Differences in DOC composition and quality were characterized with fluorescence spectroscopy. Rapidly decomposing leaves with lower tannin and lignin concentrations leached ∼40 to 50% more DOC and total dissolved N than did slowly decomposing leaves. Rates of heterotrophic respiration were 25 to 50% higher on leachate from rapidly decomposing leaf types. Rates of heterotrophic respiration were related to metrics of aromaticity. Specifically, rates of respiration were correlated negatively with the Fluorescence Index and positively with Specific Ultraviolet Absorbance (SUVA254) and T280 tryptophan-like fluorescence peak. These results reveal that leaf-litter DOC is distinctly different from ambient streamwater DOC. The relationships between optical characteristics of leaf leachate and bioavailability are opposite those found in streamwater DOC. Differences in phytochemistry among leaf types can influence stream ecosystems with respect to DOC quantity, composition, and rates of stream respiration. These patterns suggest that the relationship between the chemical structure of DOC and its biogeochemistry is more complex than previously recognized. These unique properties of leaflitter DOC will be important when assessing the effects of terrestrial C on aquatic ecosystems, especially during leaf fall.
2014
Balvanera P, Siddique I, Dee L, Paquette A, Isbell F, Gonzalez A, Byrnes J, O’Connor MI, Hungate BA, Griffin JN (2014) Linking Biodiversity and Ecosystem Services: Current Uncertainties and the Necessary Next Steps. BioScience 64(1): 49-57.
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Read PublicationUnderstanding when biodiversity conservation and ecosystem-service maintenance are compatible is needed within the Intergovernmental Platform on Biodiversity and Ecosystem Services (IPBES). Here, we evaluate current understanding and uncertainties of the effects of biodiversity change on selected ecosystem services and suggest ways to further understand the links between biodiversity change and ecosystem services. We reviewed experiments, observations, and syntheses on the effects of species richness on six ecosystem services: forage, timber, fisheries, climate regulation, agricultural pest control, and water quality. Establishing a direct link from biodiversity to ecosystem-service provision has often been precluded by limited data (i.e., the amount, consistency, or generality of the data) and a mismatch between the variables measured and the final ecosystem service that is relevant to stakeholders. We suggest that encompassing syntheses and a network of interdisciplinary experiments under realistic conditions could fill these gaps and could inform the outcomes of alternative management and policy scenarios within IPBES.
Barron-Gafford GA, Cable JM, Bentley LP, Scott RL, Huxman TE, Jenerette GD, Ogle K (2014) Quantifying the timescales over which exogenous and endogenous conditions affect soil respiration. New Phytologist 202(2): 442-454.
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Read PublicationUnderstanding how exogenous and endogenous factors and above-ground-below-ground linkages modulate carbon dynamics is difficult because of the influences of antecedent conditions. For example, there are variable lags between above-ground assimilation and below-ground efflux, and the duration of antecedent periods are often arbitrarily assigned. Nonetheless, developing models linking above- and below-ground processes is crucial for estimating current and future carbon dynamics. We collected data on leaf-level photosynthesis (Asat ) and soil respiration (Rsoil ) in different microhabitats (under shrubs vs under bunchgrasses) in the Sonoran Desert. We evaluated timescales over which endogenous and exogenous factors control Rsoil by analyzing data in the context of a semimechanistic temperature-response model of Rsoil that incorporated effects of antecedent exogenous (soil water) and endogenous (Asat ) conditions. For both microhabitats, antecedent soil water and Asat significantly affected Rsoil , but Rsoil under shrubs was more sensitive to Asat than that under bunchgrasses. Photosynthetic rates 1 and 3 d before the Rsoil measurement were most important in determining current-day Rsoil under bunchgrasses and shrubs, respectively, indicating a significant lag effect. Endogenous and exogenous controls are critical drivers of Rsoil , but the relative importance and the timescale over which each factor affects Rsoil depends on above-ground vegetation and ecosystem structure characteristics.
Briggs BR, Brodie EL, Tom LM, Dong H, Jiang H, Huang Q, Wang S, Hou W, Wu G, Huang L, Hedlund BP, Zhang C, Dijkstra P, Hungate BA (2014) Seasonal patterns in microbial communities inhabiting the hot springs of Tengchong, Yunnan Province, China. Environmental Microbiology 16(6): 1579-1591.
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Read PublicationStudies focusing on seasonal dynamics of microbial communities in terrestrial and marine environments are common; however, little is known about seasonal dynamics in high-temperature environments. Thus, our objective was to document the seasonal dynamics of both the physicochemical conditions and the microbial communities inhabiting hot springs in Tengchong County, Yunnan Province, China. The PhyloChip microarray detected 4882 operational taxonomic units (OTUs) within 79 bacterial phylum-level groups and 113 OTUs within 20 archaeal phylum-level groups, which are additional 54 bacterial phyla and 11 archaeal phyla to those that were previously described using pyrosequencing. Monsoon samples (June 2011) showed increased concentrations of potassium, total organic carbon, ammonium, calcium, sodium and total nitrogen, and decreased ferrous iron relative to the dry season (January 2011). At the same time, the highly ordered microbial communities present in January gave way to poorly ordered communities in June, characterized by higher richness of <em>B</em><em>acteria</em>, including microbes related to mesophiles. These seasonal changes in geochemistry and community structure are likely due to high rainfall influx during the monsoon season and indicate that seasonal dynamics occurs in high-temperature environments experiencing significant changes in seasonal recharge. Thus, geothermal environments are not isolated from the surrounding environment and seasonality affects microbial ecology.
Evans RD, Koyama A, Sonderegger DL, Charlet TN, Newingham BA, Fenstermaker LF, Harlow B, Jin VL, Ogle K, Smith SD, Nowak RS (2014) Greater ecosystem carbon in the Mojave Desert after ten years exposure to elevated CO2. Nature Climate Change 4(5): 394-397.
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Read PublicationCarbon dioxide is the main greenhouse gas inducing climate change. Increased global CO<sub><b>2</b></sub> emissions, estimated at 8.4 Pg C yr<sup>−<b>1</b></sup> at present, have accelerated from 1% yr<sup>−<b>1</b></sup>during 1990–1999 to 2.5% yr<sup>−<b>1</b></sup> during 2000–2009 (ref. <a id="ref-link-1" title="Friedlingstein, P. et al. Update on CO2 emissions. Nature Geosci. 3, 811-812 (2010)." href="http://www.nature.com/nclimate/journal/v4/n5/full/nclimate2184.html#ref1">1</a>). The carbon balance of terrestrial ecosystems is the greatest unknown in the global C budget because the actual magnitude, location and causes of terrestrial sinks are uncertain<sup><a id="ref-link-2" title="Ballantyne, A. P., Alden, C. B., Miller, J. B., Tans, P. P. & White, J. W. C. Increase in observed net carbon dioxide uptake by land and oceans during the past 50 years. Nature 488, 70-72 (2012)." href="http://www.nature.com/nclimate/journal/v4/n5/full/nclimate2184.html#ref2">2</a></sup>; estimates of terrestrial C uptake, therefore, are often based on the residuals between direct measurements of the atmospheric sink and well-constrained models of ocean uptake of CO<sub><b>2</b></sub> (ref. <a id="ref-link-3" title="Houghton, R. A., Hall, F. & Goetz, S. J. Importance of biomass in the global carbon cycle. J. Geophys. Res. 114, G00E03 (2009)." href="http://www.nature.com/nclimate/journal/v4/n5/full/nclimate2184.html#ref3">3</a>). Here we report significant terrestrial C accumulation caused by CO<sub><b>2</b></sub> enhancement to net ecosystem productivity in an intact, undisturbed arid ecosystem<sup><a id="ref-link-4" title="Billings, S. A., Schaeffer, S. M. & Evans, R. D. Trace N gas losses and N mineralization in Mojave Desert soils exposed to elevated CO2. Soil Biol. Biochem. 34, 1777-1784 (2002)." href="http://www.nature.com/nclimate/journal/v4/n5/full/nclimate2184.html#ref4">4</a>, <a id="ref-link-5" title="Housman, D. C. et al. Increases in desert shrub productivity under elevated carbon dioxide vary with water availability. Ecosystems 9, 374-385 (2006)." href="http://www.nature.com/nclimate/journal/v4/n5/full/nclimate2184.html#ref5">5</a>, <a id="ref-link-6" title="Ferguson, S. D. & Nowak, R. S. Transitory effects of elevated atmospheric CO2 on fine root dynamics in an arid ecosystem do not increase long-term soil carbon input from fine root litter. New Phytol. 190, 953-967 (2011)." href="http://www.nature.com/nclimate/journal/v4/n5/full/nclimate2184.html#ref6">6</a>, <a id="ref-link-7" title="Billings, S. A., Schaeffer, S. M. & Evans, R. D. Soil microbial activity and N availability with elevated CO2 in Mojave Desert soils. Glob. Biogeochem. Cycles 18, GB1011 (2004)." href="http://www.nature.com/nclimate/journal/v4/n5/full/nclimate2184.html#ref7">7</a>, <a id="ref-link-8" title="Jin, V. L. & Evans, R. D. Microbial 13C utilization patterns via stable isotope probing of phospholipid biomarkers in Mojave Desert soils exposed to ambient and elevated atmospheric CO2. Glob. Change Biol. 16, 2334-2344 (2010)." href="http://www.nature.com/nclimate/journal/v4/n5/full/nclimate2184.html#ref8">8</a></sup> following ten years of exposure to elevated atmospheric CO<sub><b>2</b></sub>. Results provide direct evidence that CO<sub><b>2</b></sub>fertilization substantially increases ecosystem C storage and that arid ecosystems are significant, previously unrecognized, sinks for atmospheric CO<sub><b>2</b></sub> that must be accounted for in efforts to constrain terrestrial and global C cycles.
Flores GE, Caporaso JG, Henley JB, Rideout JR, Domogala D, Chase J, Leff J, Vazquez Baeza Y, Gonzalez A, Knight R, Dunn RR, Fierer N (2014) Temporal variability is a personalized feature of the human microbiome. Genome Biology 15(12): 531.
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<h3>Background</h3>
<p id="__p1" class="p p-first-last">It is now apparent that the complex microbial communities found on and in the human body vary across individuals. What has largely been missing from previous studies is an understanding of how these communities vary over time within individuals. To the extent to which it has been considered, it is often assumed that temporal variability is negligible for healthy adults. Here we address this gap in understanding by profiling the forehead, gut (fecal), palm, and tongue microbial communities in 85 adults, weekly over 3 months.</p>
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<h3>Results</h3>
<p id="__p2" class="p p-first-last">We found that skin (forehead and palm) varied most in the number of taxa present, whereas gut and tongue communities varied more in the relative abundances of taxa. Within each body habitat, there was a wide range of temporal variability across the study population, with some individuals harboring more variable communities than others. The best predictor of these differences in variability across individuals was microbial diversity; individuals with more diverse gut or tongue communities were more stable in composition than individuals with less diverse communities.</p>
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<h3>Conclusions</h3>
<p id="__p3" class="p p-first-last">Longitudinal sampling of a relatively large number of individuals allowed us to observe high levels of temporal variability in both diversity and community structure in all body habitats studied. These findings suggest that temporal dynamics may need to be considered when attempting to link changes in microbiome structure to changes in health status. Furthermore, our findings show that, not only is the composition of an individual’s microbiome highly personalized, but their degree of temporal variability is also a personalized feature.</p>
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Hagerty SB, van Groenigen KJ, Allison SD, Hungate BA, Schwartz E, Koch GW, Kolka RK, Dijkstra P (2014) Accelerated microbial turnover but constant growth efficiency with warming in soil. Nature Climate Change 4(10): 903-906.
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Read PublicationRising temperatures are expected to reduce global soil carbon (C) stocks, driving a positive feedback to climate change<sup><a id="ref-link-1" title="Davidson, E. A. & Janssens, I. A. Temperature sensitivity of soil carbon decomposition and feedbacks to climate change. Nature 440, 165-173 (2006)." href="http://www.nature.com/nclimate/journal/v4/n10/full/nclimate2361.html#ref1">1</a>, <a id="ref-link-2" title="Cox, P. M., Betts, R. A., Jones, C. D., Spall, S. A. & Totterdell, I. J. Acceleration of global warming due to carbon-cycle feedbacks in a coupled climate model. Nature 408, 184-187 (2000)." href="http://www.nature.com/nclimate/journal/v4/n10/full/nclimate2361.html#ref2">2</a>, <a id="ref-link-3" title="Schlesinger, W. H. & Andrews, J. A. Soil respiration and the global carbon cycle. Biogeochemistry 48, 7-20 (2000)." href="http://www.nature.com/nclimate/journal/v4/n10/full/nclimate2361.html#ref3">3</a></sup>. However, the mechanisms underlying this prediction are not well understood, including how temperature affects microbial enzyme kinetics, growth efficiency (MGE), and turnover<sup><a id="ref-link-4" title="Agren, G. I. & Wetterstedt, J. A. M. What determines the temperature response of soil organic matter decomposition? Soil Biol. Biochem. 39, 1794-1798 (2007)." href="http://www.nature.com/nclimate/journal/v4/n10/full/nclimate2361.html#ref4">4</a>, <a id="ref-link-5" title="Li, J. W., Wang, G. S., Allison, S. D., Mayes, M. A. & Luo, Y. Q. Soil carbon sensitivity to temperature and carbon use efficiency compared across microbial-ecosystem models of varying complexity. Biogeochemistry 119, 67-84 (2014)." href="http://www.nature.com/nclimate/journal/v4/n10/full/nclimate2361.html#ref5">5</a></sup>. Here, in a laboratory study, we show that microbial turnover accelerates with warming and, along with enzyme kinetics, determines the response of microbial respiration to temperature change. In contrast, MGE, which is generally thought to decline with warming<sup><a id="ref-link-6" title="Allison, S. D., Wallenstein, M. D. & Bradford, M. A. Soil-carbon response to warming dependent on microbial physiology. Nature Geosci. 3, 336-340 (2010)." href="http://www.nature.com/nclimate/journal/v4/n10/full/nclimate2361.html#ref6">6</a>, <a id="ref-link-7" title="Manzoni, S., Taylor, P., Richter, A., Porporato, A. & Agren, G. I. Environmental and stoichiometric controls on microbial carbon-use efficiency in soils. New Phytol. 196, 79-91 (2012)." href="http://www.nature.com/nclimate/journal/v4/n10/full/nclimate2361.html#ref7">7</a>, <a id="ref-link-8" title="Sinsabaugh, R. L., Manzoni, S., Moorhead, D. L. & Richter, A. Carbon use efficiency of microbial communities: Stoichiometry, methodology and modelling. Ecol. Lett. 16, 930-939 (2013)." href="http://www.nature.com/nclimate/journal/v4/n10/full/nclimate2361.html#ref8">8</a></sup>, showed no temperature sensitivity. A microbial-enzyme model suggests that such temperature sensitive microbial turnover would promote soil C accumulation with warming, in contrast to reduced soil C predicted by traditional biogeochemical models. Furthermore, the effect of increased microbial turnover differs from the effects of reduced MGE, causing larger increases in soil C stocks. Our results demonstrate that the response of soil C to warming is affected by changes in microbial turnover. This control should be included in the next generation of models to improve prediction of soil C feedbacks to warming.
Hugelius G, Strauss J, Zubrzycki S, Harden JW, Schuur EAG, Ping CL, Schirrmeister, L, Grosse G, Michaelson GJ, Koven CD, O'Donnell JA, Elberling B, Mishra U, Camill P, Yu Z, Palmtag J, Kuhry P (2014) Estimated stocks of circumpolar permafrost carbon with quantified uncertainty ranges and identified data gaps. Biogeosciences 11(23): 6573-6593.
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Read PublicationSoils and other unconsolidated deposits in the northern circumpolar permafrost region store large amounts of soil organic carbon (SOC). This SOC is potentially vulnerable to remobilization following soil warming and permafrost thaw, but SOC stock estimates were poorly constrained and quantitative error estimates were lacking. This study presents revised estimates of permafrost SOC stocks, including quantitative uncertainty estimates, in the 0–3 m depth range in soils as well as for sediments deeper than 3 m in deltaic deposits of major rivers and in the Yedoma region of Siberia and Alaska. Revised estimates are based on significantly larger databases compared to previous studies. Despite this there is evidence of significant remaining regional data gaps. Estimates remain particularly poorly constrained for soils in the High Arctic region and physiographic regions with thin sedimentary overburden (mountains, highlands and plateaus) as well as for deposits below 3 m depth in deltas and the Yedoma region. While some components of the revised SOC stocks are similar in magnitude to those previously reported for this region, there are substantial differences in other components, including the fraction of perennially frozen SOC. Upscaled based on regional soil maps, estimated permafrost region SOC stocks are 217 ± 12 and 472 ± 27 Pg for the 0–0.3 and 0–1 m soil depths, respectively (±95% confidence intervals). Storage of SOC in 0–3 m of soils is estimated to 1035 ± 150 Pg. Of this, 34 ± 16 Pg C is stored in poorly developed soils of the High Arctic. Based on generalized calculations, storage of SOC below 3 m of surface soils in deltaic alluvium of major Arctic rivers is estimated as 91 ± 52 Pg. In the Yedoma region, estimated SOC stocks below 3 m depth are 181 ± 54 Pg, of which 74 ± 20 Pg is stored in intact Yedoma (late Pleistocene ice- and organic-rich silty sediments) with the remainder in refrozen thermokarst deposits. Total estimated SOC storage for the permafrost region is ∼1300 Pg with an uncertainty range of ∼1100 to 1500 Pg. Of this, ∼500 Pg is in non-permafrost soils, seasonally thawed in the active layer or in deeper taliks, while ∼800 Pg is perennially frozen. This represents a substantial ∼300 Pg lowering of the estimated perennially frozen SOC stock compared to previous estimates.
Hungate BA, Duval BD, Dijkstra P, Johnson DW, Ketterer ME, Stiling P, Cheng W, Millman J, Hartley A, Stover DB (2014) Nitrogen inputs and losses in response to chronic CO2 exposure in a subtropical oak woodland. Biogeosciences 11(12): 3323-3337.
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Read PublicationRising atmospheric CO2 concentrations may alter the nitrogen (N) content of ecosystems by changing N inputs and N losses, but responses vary in field experiments, possibly because multiple mechanisms are at play. We measured N fixation and N losses in a subtropical oak woodland exposed to 11 years of elevated atmospheric CO2 concentrations. We also explored the role of herbivory, carbon limitation, and competition for light or nutrients in shaping the response of N fixation to elevated CO2. Elevated CO2 did not significantly alter gaseous N losses, but lower recovery and deeper distribution in the soil of a long-term 15N tracer indicated that elevated CO2 increased leaching losses. Elevated CO2 had no effect on nonsymbiotic N fixation, and had a transient effect on symbiotic N fixation by the dominant legume. Elevated CO2 tended to reduce soil and plant concentrations of iron, molybdenum, phosphorus, and vanadium, nutrients essential for N fixation. Competition for nutrients and herbivory likely contributed to the declining response of N fixation to elevated CO2. These results indicate that positive responses of N fixation to elevated CO2 may be transient and that chronic exposure to elevated CO2 can increase N leaching. Models that assume increased fixation or reduced N losses with elevated CO2 may overestimate future N accumulation in the biosphere.
Hurteau MD, Robards TA, Stevens D, Saah D, North M, Koch GW (2014) Modeling climate and fuel reduction impacts on mixed-conifer forest carbon stocks in the Sierra Nevada, California. Forest Ecology and Management 315: 30-42.
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Read PublicationQuantifying the impacts of changing climatic conditions on forest growth is integral to estimating future forest carbon balance. We used a growth-and-yield model, modified for climate sensitivity, to quantify the effects of altered climate on mixed-conifer forest growth in the Lake Tahoe Basin, California. Estimates of forest growth and live tree carbon stocks were made for low and high emission scenarios using four downscaled general circulation model (GCM) projections. The climate scenarios were coupled with a range of commonly-used fuels reduction treatments to quantify the combined effects of these factors on live tree carbon stocks. We compared mid- (2020–2049) and late-21st (2070–2099) century carbon stock estimates with a baseline period of 1970–1999 using common input data across time periods. Recursive partitioning analysis indicates that GCM, forest composition, and simulation period most influence live tree carbon stock changes. Comparison with the late 20th century baseline period shows mixed carbon stock responses across scenarios. Growth varied by species, often with compensatory responses among dominant species that limited changes in total live tree carbon. The influence of wildfire mitigation treatments was relatively consistent with each GCM by emission scenario combination. Treatments that included prescribed fire had greater live tree carbon gains relative to baseline under the scenarios that had overall live tree carbon gains. However, across GCMs the influence of treatments varied considerably among GCM projections, indicating that further refinement of regional climate projections will be required to improve model estimates of fuel manipulations on forest carbon stocks. Additionally, had out simulations included the effects of projected climate changes on increasing wildfire probability, the effects of management treatments on carbon stocks may have been more pronounced because of the influence of treatment on fire severity.
Koch BJ, Febria CM, Gevrey M, Wainger LA, Palmer MA (2014) Nitrogen Removal by Stormwater Management Structures: A Data Synthesis. JAWRA Journal of the American Water Resources Association 50(6): 1594-1607.
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Read PublicationA comprehensive synthesis of data from empirically based published studies and a widely used stormwater best management practice (BMP) database were used to assess the variability in nitrogen (N) removal performance of urban stormwater ponds, wetlands, and swales and to identify factors that may explain this variability. While the data suggest that BMPs were generally effective on average, removal efficiencies of ammonium (NH<sub>4</sub>), nitrate (NO<sub>3</sub>), and total nitrogen (TN) were highly variable ranging from negative (i.e., BMPs acting as sources of N) to 100%. For example, removal of NO<sub>3</sub> varied from (median ±1 SD) −15 ± 49% for dry ponds, 32 ± 120% for wet ponds, 58 ± 210% for wetlands, and 37 ± 29% for swales. Across the same BMP types, TN removal was 27 ± 24%, 40 ± 31%, 61 ± 30%, and 50 ± 29%. NH<sub>4</sub> removal was 9 ± 36%, 29 ± 72%, 31 ± 24%, and 45 ± 34%. BMP size, age, and location explained some of the variability. For example, small and shallow ponds and wetlands were more effective than larger, deeper ones in removing N. Despite well-known intra-annual variation in N fluxes, most measurements have been made over short time periods using concentrations, not flow-weighted N fluxes. Urban N export is increasing in some areas as large storms become more frequent. Thus, accounting for the full range of BMP performance under such conditions is crucial. A select number of long-term flux-based BMP studies that rigorously measure rainfall, hydrology, and site conditions could improve BMP implementation.
Langley JA, Hungate BA (2014) Plant community feedbacks and long-term ecosystem responses to multi-factored global change. AoB Plants (6): plu035.
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Read PublicationWhile short-term plant responses to global change are driven by physiological mechanisms, which are represented relatively well by models, long-term ecosystem responses to global change may be determined by shifts in plant community structure resulting from other ecological phenomena such as interspecific interactions, which are represented poorly by models. In single-factor scenarios, plant communities often adjust to increase ecosystem response to that factor. For instance, some early global change experiments showed that elevated CO<sub>2</sub> favours plants that respond strongly to elevated CO<sub>2</sub>, generally amplifying the response of ecosystem productivity to elevated CO<sub>2</sub>, a positive community feedback. However, most ecosystems are subject to multiple drivers of change, which can complicate the community feedback effect in ways that are more difficult to generalize. Recent studies have shown that (i) shifts in plant community structure cannot be reliably predicted from short-term plant physiological response to global change and (ii) that the ecosystem response to multi-factored change is commonly less than the sum of its parts. Here, we survey results from long-term field manipulations to examine the role community shifts may play in explaining these common findings. We use a simple model to examine the potential importance of community shifts in governing ecosystem response. Empirical evidence and the model demonstrate that with multi-factored change, the ecosystem response depends on community feedbacks, and that the magnitude of ecosystem response will depend on the relationship between plant response to one factor and plant response to another factor. Tradeoffs in the ability of plants to respond positively to, or to tolerate, different global change drivers may underlie generalizable patterns of covariance in responses to different drivers of change across plant taxa. Mechanistic understanding of these patterns will help predict the community feedbacks that determine long-term ecosystem responses.
LeRoy CJ, Wymore AS, Davis R and Marks JC (2014) Indirect influences of a major drought on leaf litter quality and decomposition in a southwestern stream. Fundamental and Applied Limnology 184(1): 1–10.
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Read PublicationClimate models predict that the southwestern United States will experience an increase in drought frequency and intensity with global climate change. We tested the hypothesis that leaf litter produced under natural drought conditions would have an altered litter chemistry profile and affect decomposition rates and macro invertebrate colonization compared to non-drought conditions. To test this hypothesis we collected leaf litter from Populus fremontii, Alnus oblongifolia, and Platanus wrightii grown during an average precipitation year (2001) and a record drought year (2002) and performed an in-stream decomposition study using both litter types. Three major patterns emerged: 1) Drought conditions significantly altered litter chemistry for mature trees of three species; however, the direction and magnitude of change differed among species and litter chemicals; 2) Leaf litter mass loss was influenced by both differences among species and drought; yet, species effects were more pronounced over time than drought effects; and 3) After 69 days of decomposition, the structure of the macroinvertebrate community was uninfluenced by the drought effect on A. oblongifolia or P. wrightii litters, but there was a community-wide drought effect on macroinvertebrate communities colonizing P. fremontii litter. Many recent studies have explored the influence of drought on stream flow and water temperatures, but these results suggest that litter quality can change under different climatic conditions, but the overall decay of leaf material may not be dramatically altered by droughts. Understanding how forest-stream interactions may be altered by the various influences of climate change will allow for better predictions regarding how long-term disturbances may alter stream ecosystem functioning.
Li J, Luo Y, Natali S, Schuur EAG, Xia J, Kowalczyk E, Wang Y (2014) Modeling permafrost thaw and ecosystem carbon cycle under annual and seasonal warming at an Arctic tundra site in Alaska. Journal of Geophysical Research-Biogeosciences 119(6): 1129–1146.
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Read PublicationPermafrost thaw and its impacts on ecosystem carbon (C) dynamics are critical for predicting global climate change. It remains unclear whether annual and seasonal warming (winter or summer) affect permafrost thaw and ecosystem C balance differently. It is also required to compare the short-term stepwise warming and long-term gradual warming effects. This study validated a land surface model, the Community Atmosphere Biosphere Land Exchange model, at an Alaskan tundra site, and then used it to simulate permafrost thaw and ecosystem C flux under annual warming, winter warming, and summer warming. The simulations were conducted under stepwise air warming (2°C yr<sup>−1</sup>) during 2007–2011, and gradual air warming (0.04°C yr<sup>−1</sup>) during 2007–2056. We hypothesized that all warming treatments induced greater permafrost thaw, and larger ecosystem respiration than plant growth thus shifting the ecosystem C sink to C source. Results only partially supported our hypothesis. Climate warming further enhanced C sink under stepwise (6–15%) and gradual (1–8%) warming scenarios as followed by annual warming, winter warming, and summer warming. This is attributed to disproportionally low temperature increase in soil (0.1°C) in comparison to air warming (2°C). In a separate simulation, a greater soil warming (1.5°C under winter warming) led to a net ecosystem C source (i.e., 18 g C m<sup>−2</sup> yr<sup>−1</sup>). This suggests that warming tundra can potentially provide positive feedbacks to global climate change. As a key variable, soil temperature and its dynamics, especially during wintertime, need to be carefully studied under global warming using both modeling and experimental approaches.
Liu CM, Osborne BJW, Hungate BA, Shahabi K, Huibner S, Lester R, Dwan MG, Kovacs C, Contente-Cuomo TL, Benko E, Aziz M, Pricen LB, Kaul R (2014) The semen microbiome and its relationship with local immunology and viral load in HIV infection. PLOS Pathogens 10(7): e1004262.
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Read PublicationSemen is a major vector for HIV transmission, but the semen HIV RNA viral load (VL) only correlates moderately with the blood VL. Viral shedding can be enhanced by genital infections and associated inflammation, but it can also occur in the absence of classical pathogens. Thus, we hypothesized that a dysregulated semen microbiome correlates with local HIV shedding. We analyzed semen samples from 49 men who have sex with men (MSM), including 22 HIV-uninfected and 27 HIV-infected men, at baseline and after starting antiretroviral therapy (ART) using 16S rRNA gene-based pyrosequencing and quantitative PCR. We studied the relationship of semen bacteria with HIV infection, semen cytokine levels, and semen VL by linear regression, non-metric multidimensional scaling, and goodness-of-fit test.<em>Streptococcus</em>, <em>Corynebacterium</em>, and <em>Staphylococcus</em> were common semen bacteria, irrespective of HIV status. While <em>Ureaplasma</em> was the more abundant Mollicutes in HIV-uninfected men, <em>Mycoplasma</em> dominated after HIV infection. HIV infection was associated with decreased semen microbiome diversity and richness, which were restored after six months of ART. In HIV-infected men, semen bacterial load correlated with seven pro-inflammatory semen cytokines, including IL-6 (<em>p</em> = 0.024), TNF-α (<em>p</em> = 0.009), and IL-1b (<em>p</em> = 0.002). IL-1b in particular was associated with semen VL (<em>r<sup>2</sup></em> = 0.18, <em>p</em> = 0.02). Semen bacterial load was also directly linked to the semen HIV VL <em>(r<sup>2</sup></em> = 0.15, <em>p</em> = 0.02). HIV infection reshapes the relationship between semen bacteria and pro-inflammatory cytokines, and both are linked to semen VL, which supports a role of the semen microbiome in HIV sexual transmission.
Liu XJA, Fike JH, Galbraith JM, Fike WB (2014) Switchgrass Response to Cutting Frequency and Biosolids Amendment: Biomass Yield, Feedstock Quality, and Theoretical Ethanol Yield. BioEnergy Research 7(4): 1191-1200.
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Read PublicationBiofuel crops have relatively low economic value, and potential to grow them with low-cost inputs is essential for economic viability. Use of biosolids as a fertility source has not been explored at the field scale for switchgrass (<em class="EmphasisTypeItalic ">Panicum virgatum</em> L.), a potential bioenergy crop. This study tested harvest management and biosolids rate effects on switchgrass production, quality, and theoretical ethanol yield in Virginia, USA. Switchgrass (cv. “Cave-in-Rock”) was annually cut once (winter) or twice (summer and winter) for 2 years. Biosolids were applied once at 0, 77, and 154 kg N ha<sup>−1</sup> in May 2011; urea was applied once at 146 kg N ha<sup>−1</sup> for comparison. Feedstock yield and quality parameters (neutral and acid detergent fibers, cellulose, hemicellulose, lignin, and ash) were measured and used to compute theoretical ethanol potential (TEP) and theoretical ethanol yield (TEY). Cutting twice per season produced greater biomass yields than cutting once (6.6 vs 5.4 Mg ha<sup>−1</sup>) in 2011 but not in 2012. Cutting once per season produced feedstock with greater TEP (513 vs 433 L Mg<sup>−1</sup>) and TEY (2,980 vs 2,680 L ha<sup>−1</sup>) in both years. Biosolids and urea increased biomass yields by 11 % (0.6 Mg ha<sup>−1</sup>) and TEY by 13 % (352 L Mg<sup>−1</sup>), but both decreased TEP by 1 % (7.1 L Mg<sup>−1</sup> biomass). Cutting once per season is advantageous in producing more TEY given comparable biomass yield and superior feedstock quality. Biosolids were a suitable alternate N source and could boost biomass and biofuel production while reducing input costs in switchgrass-based bioenergy systems.
Luo C, Rodriguez-R LM, Johnston ER, Wu L, Cheng L, Xue K, Tu Q, Deng Y, He Z, Shi JZ, Yuan MM, Sherry RA, Li D, Luo Y, Schuur EAG, Chain P, Tiedje JM, Zhou J, Konstantinidis, KT (2014) Soil Microbial Community Responses to a Decade of Warming as Revealed by Comparative Metagenomics. Applied and Environmental Microbiology 80(5): 1777-1786.
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Read PublicationSoil microbial communities are extremely complex, being composed of thousands of low-abundance species (<0.1% of total). How such complex communities respond to natural or human-induced fluctuations, including major perturbations such as global climate change, remains poorly understood, severely limiting our predictive ability for soil ecosystem functioning and resilience. In this study, we compared 12 whole-community shotgun metagenomic data sets from a grassland soil in the Midwestern United States, half representing soil that had undergone infrared warming by 2°C for 10 years, which simulated the effects of climate change, and the other half representing the adjacent soil that received no warming and thus, served as controls. Our analyses revealed that the heated communities showed significant shifts in composition and predicted metabolism, and these shifts were community wide as opposed to being attributable to a few taxa. Key metabolic pathways related to carbon turnover, such as cellulose degradation (∼13%) and CO<sub>2</sub> production (∼10%), and to nitrogen cycling, including denitrification (∼12%), were enriched under warming, which was consistent with independent physicochemical measurements. These community shifts were interlinked, in part, with higher primary productivity of the aboveground plant communities stimulated by warming, revealing that most of the additional, plant-derived soil carbon was likely respired by microbial activity. Warming also enriched for a higher abundance of sporulation genes and genomes with higher G+C content. Collectively, our results indicate that microbial communities of temperate grassland soils play important roles in mediating feedback responses to climate change and advance the understanding of the molecular mechanisms of community adaptation to environmental perturbations.
Mauritz M, Cleland E, Merkley M, Lipson DA (2014) The Influence of Altered Rainfall Regimes on Early Season N Partitioning Among Early Phenology Annual Plants, a Late Phenology Shrub, and Microbes in a Semi-arid Ecosystem. Ecosystems 17(8): 1354-1370.
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Read PublicationIn Mediterranean-type ecosystems, nitrogen (N) accumulates in soil during dry summer months and rapidly becomes available during early season rain events. The availability of early season N could depend on the size of rainfall events, soil microbial activity, and phenology of the plant community. However, it is poorly understood how precipitation patterns affect the fate of early season N. Microbes and plants with early phenology may compete strongly for early season N but theory suggests that microbial N storage can meet plant N demands later in the season. Using a <sup>15</sup>N tracer and rainfall manipulation we investigated the fate of early season N. N allocation patterns differed substantially between microbes, early and late phenology plants. As expected early phenology annuals and microbes took up <sup>15</sup>N, within 1 day, whereas a late-phenology shrub allocated <sup>15</sup>N to leaves later in the season. We saw no evidence for microbial storage of early season N; the peak of <sup>15</sup>N in shrub leaves did not coincide with detectable levels of <sup>15</sup>N in the microbial biomass or labile soil pool. This suggests that shrubs were able to access early season N, store and allocate it for growth later in the season. Although we saw no evidence of microbial N storage, N retention in soil organic matter (SOM) was high and microbes may play an important role in sequestering N to SOM. Plant N uptake did not respond significantly to 1 year of rainfall manipulation, but microbes were sensitive to dry conditions. 1 year after <sup>15</sup>N addition shrubs had resorbed up to half of the N from leaves whereas N in annuals remained as dead leaf litter. Differences in end-of-season N partitioning between dead and living biomass in the two vegetation types suggest that plant species composition could affect N availability in the following growing season, but it may take several years of altered precipitation patterns to produce rainfall-dependent changes.
Mayor JR, Wright SJ, Schuur EAG, Brooks ME, Turner BL (2014) Stable nitrogen isotope patterns of trees and soils altered by long-term nitrogen and phosphorus addition to a lowland tropical rainforest. Biogeochemistry 119(1): 293-306.
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Read PublicationFoliar nitrogen (N) isotope ratios (δ<sup>15</sup>N) are used as a proxy for N-cycling processes, including the “openness” of the N cycle and the use of distinct N sources, but there is little experimental support for such proxies in lowland tropical forest. To address this, we examined the δ<sup>15</sup>N values of soluble soil N and canopy foliage of four tree species after 13 years of factorial N and P addition to a mature lowland rainforest. We hypothesized that N addition would lead to <sup>15</sup>N-enriched soil N forms due to fractionating losses, whereas P addition would reduce N losses as the plants and microbes adjusted their stoichiometric demands. Chronic N addition increased the concentration and δ<sup>15</sup>N value of soil nitrate and δ<sup>15</sup>N in live and senesced leaves in two of four tree species, but did not affect ammonium or dissolved organic N. Phosphorus addition significantly increased foliar δ<sup>15</sup>N in one tree species and elicited significant N × P interactions in two others due to a reduction in foliar δ<sup>15</sup>N enrichment under N and P co-addition. Isotope mixing models indicated that three of four tree species increased their use of nitrate relative to ammonium following N addition, supporting the expectation that tropical trees use the most available form of mineral N. Previous observations that anthropogenic N deposition in this tropical region have led to increasing foliar δ<sup>15</sup>N values over decadal time-scales is now mechanistically linked to greater usage of <sup>15</sup>N-enriched nitrate.
McHugh T, Schwartz E, Koch G, Pett-Ridge J, Weber P, Hungate B (2014) Multiple-Element Isotope Probes, NanoSIMS, and the Functional Genomics of Microbial Carbon Cycling in Soils in Response to Chronic Climatic Change. 2014 Genomic Science Contractor-Grantee Meeting XII 93.
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Read PublicationFor the past several decades, connecting biogeochemistry and microbial genomics has been a high priority in microbial ecology. Yet, techniques that actually link element flow and genomic information are scarce. In this project, we are using the Chip-SIP method to measure isotopic composition of major elements (C, N, H, and O) of nucleic acid sequences representing individual microbial taxa. RNA is extracted from an environmental sample after exposure to isotopically labeled substrates. The nucleic acids from the entire microbial community are then exposed to a microarray containing small probes that target the 16S rRNA genes of a large variety of microorganisms so that nucleic acids extracted from the environmental sample bind to matching probes. Then, the entire microarray is placed under a nanoscale secondary ion mass spectrometer, which sequentially analyzes the RNA bound to each probe for isotopic composition. In this way, element flow in the natural environment into individual microbial taxa can be determined.
McHugh TA, Koch GW, Schwartz E (2014) Minor Changes in Soil Bacterial and Fungal Community Composition Occur in Response to Monsoon Precipitation in a Semiarid Grassland. Microbial Ecology 68(2): 370-378.
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Read PublicationArizona and New Mexico receive half of their annual precipitation during the summer monsoon season, making this large-scale rain event critical for ecosystem productivity. We used the monsoon rains to explore the responses of soil bacterial and fungal communities to natural moisture pulses in a semiarid grassland. Through 454 pyrosequencing of the 16S rRNA gene and ITS region, we phylogenetically characterized these communities at 22 time points during a summer season. Relative humidity increased before the rains arrived, creating conditions in soil that allowed for the growth of microorganisms. During the course of the study, the relative abundances of most bacterial phyla showed little variation, though some bacterial populations responded immediately to an increase in soil moisture once the monsoon rains arrived. The Firmicutes phylum experienced over a sixfold increase in relative abundance with increasing water availability. Conversely, Actinobacteria, the dominant taxa at our site, were negatively affected by the increase in water availability. No relationship was found between bacterial diversity and soil water potential. Bacterial community structure was unrelated to all environmental variables that we measured, with the exception of a significant relationship with atmospheric relative humidity. Relative abundances of fungal phyla fluctuated more throughout the season than bacterial abundances did. Variation in fungal community structure was unrelated to soil water potential and to most environmental variables. However, ordination analysis showed a distinct fungal community structure late in the season, probably due to plant senescence.
Morrissey EM, Berrier DJ, Neubauer SC, Franklin RB (2014) Using microbial communities and extracellular enzymes to link soil organic matter characteristics to greenhouse gas production in a tidal freshwater wetland. Biogeochemistry 117(2): 473-490.
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Read PublicationTo gain a more mechanistic understanding of how soil organic matter (OM) characteristics can affect carbon mineralization in tidal freshwater wetlands, we conducted a long-term in situ field manipulation of OM type and monitored associated changes in carbon dioxide (CO<sub>2</sub>) and methane (CH<sub>4</sub>) production. In addition, we characterized microbial community structure and quantified the activity of several extracellular enzymes (EEA) involved in the acquisition of carbon, nitrogen, and phosphorus. Treatments included a plant litter addition, prepared using naturally-senescing vegetation from the site, and a compost amendment, designed to increase the concentration of aged, partially humified, OM. Both types of OM-amended soils had CO<sub>2</sub>production rates 40–50 % higher than unamended control soils, suggesting that the added OM had inherently higher quality and/or availability than the native soil OM. Rates of CO<sub>2</sub> production were not correlated with microbial community structure or EEA except a modest relationship with cellulose breakdown via the K<sub>m</sub> of β-1,4-glucosidase. We interpret this lack of correlation to be a consequence of high functional redundancy of microorganisms that are capable of producing CO<sub>2</sub>. Rates of CH<sub>4</sub>production were also influenced by OM quality, increasing by an order of magnitude with plant litter additions relative to compost-amended and control soils. Unlike CO<sub>2</sub>, rates of CH<sub>4</sub> production were significantly correlated with the microbial community structure and with enzyme kinetic parameters (V<sub>max</sub> and K<sub>m</sub>) for both carbon (β-1,4-glucosidase, 1,4-β-cellobiosidase, and β-<span class="EmphasisTypeSmallCaps ">d</span>-xylosidase) and nitrogen acquisition (leucyl aminopeptidase). The monophyletic nature of methanogenic archaea, combined with their reliance on a small select group of organic substrates produced via enzyme-mediated hydrolysis and subsequent bacterial fermentation, provides a basis for the strong links between microbial community structure, EEA, and CH<sub>4</sub> production. Our results suggest that incorporating microbial community structure and EEA into conceptual models of wetland OM decomposition may enhance our mechanistic understanding of, and predictive capacity for, biogeochemical process rates.
Morrissey EM, Gillespie JL, Morina JC, Franklin RB (2014) Salinity affects microbial activity and soil organic matter content in tidal wetlands. Global Change Biology 20(4): 1351-1362.
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Read PublicationClimate change-associated sea level rise is expected to cause saltwater intrusion into many historically freshwater ecosystems. Of particular concern are tidal freshwater wetlands, which perform several important ecological functions including carbon sequestration. To predict the impact of saltwater intrusion in these environments, we must first gain a better understanding of how salinity regulates decomposition in natural systems. This study sampled eight tidal wetlands ranging from freshwater to oligohaline (0–2 ppt) in four rivers near the Chesapeake Bay (Virginia). To help isolate salinity effects, sites were selected to be highly similar in terms of plant community composition and tidal influence. Overall, salinity was found to be strongly negatively correlated with soil organic matter content (OM%) and C : N, but unrelated to the other studied environmental parameters (pH, redox, and above- and below-ground plant biomass). Partial correlation analysis, controlling for these environmental covariates, supported direct effects of salinity on the activity of carbon-degrading extracellular enzymes (β-1, 4-glucosidase, 1, 4-β-cellobiosidase, β-D-xylosidase, and phenol oxidase) as well as alkaline phosphatase, using a per unit OM basis. As enzyme activity is the putative rate-limiting step in decomposition, enhanced activity due to salinity increases could dramatically affect soil OM accumulation. Salinity was also found to be positively related to bacterial abundance (qPCR of the 16S <em>rRNA</em> gene) and tightly linked with community composition (T-RFLP). Furthermore, strong relationships were found between bacterial abundance and/or composition with the activity of specific enzymes (1, 4-β-cellobiosidase, arylsulfatase, alkaline phosphatase, and phenol oxidase) suggesting salinity's impact on decomposition could be due, at least in part, to its effect on the bacterial community. Together, these results indicate that salinity increases microbial decomposition rates in low salinity wetlands, and suggests that these ecosystems may experience decreased soil OM accumulation, accretion, and carbon sequestration rates even with modest levels of saltwater intrusion.
Natali SM, Schuur EAG, Webb E, Pries E, Hicks CE, Crummer KG (2014) Permafrost degradation stimulates carbon loss from experimentally warmed tundra. Ecology 95(3): 602-608.
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Read PublicationA large pool of organic carbon (C) has been accumulating in the Arctic for thousands of years because cold and waterlogged conditions have protected soil organic material from microbial decomposition. As the climate warms this vast and frozen C pool is at risk of being thawed, decomposed, and released to the atmosphere as greenhouse gasses. At the same time, some C losses may be offset by warming-mediated increases in plant productivity. Plant and microbial responses to warming ultimately determine net C exchange from ecosystems, but the timing and magnitude of these responses remain uncertain. Here we show that experimental warming and permafrost (ground that remains below 0°C for two or more consecutive years) degradation led to a two-fold increase in net ecosystem C uptake during the growing season. However, warming also enhanced winter respiration, which entirely offset growing-season C gains. Winter C losses may be even higher in response to actual climate warming than to our experimental manipulations, and, in that scenario, could be expected to more than double overall net C losses from tundra to the atmosphere. Our results highlight the importance of winter processes in determining whether tundra acts as a C source or sink, and demonstrate the potential magnitude of C release from the permafrost zone that might be expected in a warmer climate.
Ogle K, Pathikonda S, Sartor K, Lichstein JW, Osnas JLD, Pacala SW (2014) A model-based meta-analysis for estimating species-specific wood density and identifying potential sources of variation. Journal of Ecology 102(1): 194-208.
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Read PublicationPlant functional traits are important determinants of survival and fitness, and wood density (WD) is a key trait linked to mechanical stability, growth rates and drought- and shade-tolerance strategies. Thus, rigorous WD estimates are necessary to identify factors affecting tree performance. We obtained 1766 records of WD from the literature for 141 tree species in the United States. We implemented a hierarchical Bayesian (HB) meta-analysis that incorporated sample size, variance, covariate (e.g. moisture content and latewood proportion) and methodological information to obtain standardized estimates of WD for 305 U.S. tree species. The HB framework allowed ’borrowing of strength’ between species such that WD estimates for data-poor species were informed by data-rich species via taxonomic or phylogenetic relationships. After accounting for important covariates and sampling effects, evaluation of the residual variation revealed the potential importance of environmental factors and evolutionary history. Differential variation in WD between species within genera and between genera within orders suggested that WD is relatively conserved in some genera and orders, but not in others. WD also varied between studies (or sites) indicating the potential influence of edaphic, topographic, or population factors on intraspecific variation in WD. Synthesis. Our hierarchical Bayesian approach overcomes many of the limitations of traditional meta-analyses, and the incorporation of phylogenetic or taxonomic information facilitates estimates of trait values for data-poor species. We provide relatively well-constrained WD estimates for 305 tree species, which may be useful for tree growth and forest models, and the uncertainties associated with the estimates may inform future sampling campaigns.
Palmer MA, Hondula KL, Koch BJ (2014) Ecological Restoration of Streams and Rivers: Shifting Strategies and Shifting Goals. Annual Review of Ecology, Evolution, and Systematics 45(1): 247-269.
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Read PublicationEcological restoration has grown rapidly and now encompasses not only classic ecological theory but also utilitarian concerns, such as preparedness for climate change and provisioning of ecosystem services. Three dominant perspectives compete to influence the science and practice of river restoration. A strong focus on channel morphology has led to approaches that involve major Earth-moving activities, such as channel reconfiguration with the unmet assumption that ecological recovery will follow. Functional perspectives of river restoration aim to regain the full suite of biogeochemical, ecological, and hydrogeomorphic processes that make up a healthy river, and though there is well-accepted theory to support this, research on methods to implement and assess functional restoration projects is in its infancy. A plethora of new studies worldwide provide data on why and how rivers are being restored as well as the project outcomes. Measurable improvements postrestoration vary by restoration method and measure of outcome.
Pastor A, Compson ZG, Dijkstra P, Riera JL, Martí E, Sabater F, Hungate BA, Marks JC (2014) Stream carbon and nitrogen supplements during leaf litter decomposition: contrasting patterns for two foundation species. Oecologia 176(3): 1111-1121.
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Read PublicationLeaf litter decomposition plays a major role in nutrient dynamics in forested streams. The chemical composition of litter affects its processing by microorganisms, which obtain nutrients from litter and from the water column. The balance of these fluxes is not well known, because they occur simultaneously and thus are difficult to quantify separately. Here, we examined C and N flow from streamwater and leaf litter to microbial biofilms during decomposition. We used isotopically enriched leaves (13C and 15N) from two riparian foundation tree species: fast-decomposing Populus fremontii and slow-decomposing Populus angustifolia, which differed in their concentration of recalcitrant compounds. We adapted the isotope pool dilution method to estimate gross elemental fluxes into litter microbes. Three key findings emerged: litter type strongly affected biomass and stoichiometry of microbial assemblages growing on litter; the proportion of C and N in microorganisms derived from the streamwater, as opposed to the litter, did not differ between litter types, but increased throughout decomposition; gross immobilization of N from the streamwater was higher for P. fremontii compared to P. angustifolia, probably as a consequence of the higher microbial biomass on P. fremontii. In contrast, gross immobilization of C from the streamwater was higher for P. angustifolia, suggesting that dissolved organic C in streamwater was used as an additional energy source by microbial assemblages growing on slow-decomposing litter. These results indicate that biofilms on decomposing litter have specific element requirements driven by litter characteristics, which might have implications for whole-stream nutrient retention.
Pietsch KA, Ogle K, Cornelissen JHC, Cornwell WK, Bönisch G, Craine JM, Jackson BG, Kattge J, Peltzer DA, Penuelas J, Reich PB, Wardle DA, Weedon JT, Wright IJ, Zanne AE, Wirth C (2014) Global relationship of wood and leaf litter decomposability: the role of functional traits within and across plant organs. Global Ecology and Biogeography 23(9): 1046-1057.
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Read Publication<section id="geb12172-sec-0001" class="article-section article-body-section">
<h3>Aim</h3>
Recent meta-analyses have revealed that plant traits and their phylogenetic history influence decay rates of dead wood and leaf litter, but it remains unknown if decay rates of wood and litter covary over a wide range of tree species and across ecosystems. We evaluated the relationships between species-specific wood and leaf litter decomposability, as well as between wood and leaf traits that control their respective decomposability.
</section><section id="geb12172-sec-0002" class="article-section article-body-section">
<h3>Location</h3>
Global.
</section><section id="geb12172-sec-0003" class="article-section article-body-section">
<h3>Methods</h3>
We compiled data on rates of wood and leaf litter decomposition for 324 and 635 tree species, respectively, and data on six functional traits for both organs. We used hierarchical Bayesian meta-analysis to estimate, for the first time, species-specific values for wood and leaf litter decomposability standardized to reference conditions (<em>k</em>*<sub>wood</sub> and <em>k</em>*<sub>leaf</sub>) across the globe. With these data, we evaluated the relationships: (1) between wood and leaf traits, (2) between each <em>k</em>* and the selected traits within and across organs, and (3) between wood and leaf <em>k</em>*.
</section><section id="geb12172-sec-0004" class="article-section article-body-section">
<h3>Results</h3>
Across all species <em>k</em>*<sub>wood</sub> and <em>k</em>*<sub>leaf</sub> were positively correlated, phylogenetically clustered and correlated with plant functional traits within and across organs. <em>k</em>* of both organs was usually better described as a function of within- and cross-organ traits, than of within-organ traits alone. When analysed for angiosperms and gymnosperms separately, wood and leaf <em>k</em>* were no longer significantly correlated, but each <em>k</em>* was still significantly correlated to the functional traits.
</section><section id="geb12172-sec-0005" class="article-section article-body-section">
<h3>Main conclusions</h3>
We demonstrate important relationships among wood and leaf litter decomposability as after-life effects of traits from the living plants. These functional traits influence the decomposability of senesced tissue which could potentially lead to alterations in the rates of biogeochemical cycling, depending on the phylogenetic structure of the species pool. These results provide crucial information for a better representation of decomposition rates in dynamic global vegetation models.
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Pizano C, Baron AF, Schuur EAG, Crummer KG, Mack MC (2014) Effects of thermo-erosional disturbance on surface soil carbon and nitrogen dynamics in upland arctic tundra. Environmental Research Letters 9(7): 075006.
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Read PublicationThaw of ice-rich permafrost soils on sloping terrain can trigger erosional disturbance events that displace large volumes of soil and sediment, kill and damage plants, and initiate secondary succession. We examined how retrogressive thaw slumps (RTS), a common form of thermo-erosional disturbance in arctic tundra, affected the local loss and re-accumulation of carbon (C) and nitrogen (N) pools in organic and surface mineral soil horizons of 18 slumps within six spatially independent sites in arctic Alaska. RTS displaced 3 kg C and 0.2 kg N per m<sup>2</sup> from the soil organic horizon but did not alter pools of C and N in the top 15 cm of the mineral horizon. Surface soil C pools re-accumulated rapidly (32 ± 10 g C m<sup>−2</sup> yr<sup>−1</sup>) through the first 60 years of succession, reaching levels similar to undisturbed tundra 40–64 years after disturbance. Average N re-accumulation rates (2.2 ± 1.1 g N m<sup>−2</sup> yr<sup>−1</sup>) were much higher than expected from atmospheric deposition and biological N fixation. Finally, plant community dominance shifted from graminoids to tall deciduous shrubs, which are likely to promote higher primary productivity, biomass accumulation, and rates of nutrient cycling.
Rideout JR, He Y, Navas-Molina JA, Walters WA, Ursell LK, Gibbons SM, Chase J, McDonald D, Gonzalez A, Robbins-Pianka A, Clemente JC, Gilbert JaA, Huse SM, Zhou HW, Knight R, Caporaso JG (2014) Subsampled open-reference clustering creates consistent, comprehensive OTU definitions and scales to billions of sequences. PeerJ 2: e545.
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Read PublicationWe present a performance-optimized algorithm, subsampled open-reference OTU picking, for assigning marker gene (e.g., 16S rRNA) sequences generated on next-generation sequencing platforms to operational taxonomic units (OTUs) for microbial community analysis. This algorithm provides benefits over de novo OTU picking (clustering can be performed largely in parallel, reducing runtime) and closed-reference OTU picking (all reads are clustered, not only those that match a reference database sequence with high similarity). Because more of our algorithm can be run in parallel relative to “classic” open-reference OTU picking, it makes open-reference OTU picking tractable on massive amplicon sequence data sets (though on smaller data sets, “classic” open-reference OTU clustering is often faster). We illustrate that here by applying it to the first 15,000 samples sequenced for the Earth Microbiome Project (1.3 billion V4 16S rRNA amplicons). To the best of our knowledge, this is the largest OTU picking run ever performed, and we estimate that our new algorithm runs in less than 1/5 the time than would be required of “classic” open reference OTU picking. We show that subsampled open-reference OTU picking yields results that are highly correlated with those generated by “classic” open-reference OTU picking through comparisons on three well-studied datasets. An implementation of this algorithm is provided in the popular QIIME software package, which uses uclust for read clustering. All analyses were performed using QIIME’s uclust wrappers, though we provide details (aided by the open-source code in our GitHub repository) that will allow implementation of subsampled open-reference OTU picking independently of QIIME (e.g., in a compiled programming language, where runtimes should be further reduced). Our analyses should generalize to other implementations of these OTU picking algorithms. Finally, we present a comparison of parameter settings in QIIME’s OTU picking workflows and make recommendations on settings for these free parameters to optimize runtime without reducing the quality of the results. These optimized parameters can vastly decrease the runtime of uclust-based OTU picking in QIIME.
Schädel C, Schuur EAG, Bracho RE, Knoblauch B, Lee C, Luo H, Shaver Y, Turetsky GR, Merritt R. (2014) Circumpolar assessment of permafrost C quality and its vulnerability over time using long-term incubation data. Global Change Biology 20(2): 641-652.
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Read PublicationHigh-latitude ecosystems store approximately 1700 Pg of soil carbon (C), which is twice as much C as is currently contained in the atmosphere. Permafrost thaw and subsequent microbial decomposition of permafrost organic matter could add large amounts of C to the atmosphere, thereby influencing the global C cycle. The rates at which C is being released from the permafrost zone at different soil depths and across different physiographic regions are poorly understood but crucial in understanding future changes in permafrost C storage with climate change. We assessed the inherent decomposability of C from the permafrost zone by assembling a database of long-term (>1 year) aerobic soil incubations from 121 individual samples from 23 high-latitude ecosystems located across the northern circumpolar permafrost zone. Using a three-pool (i.e., fast, slow and passive) decomposition model, we estimated pool sizes for C fractions with different turnover times and their inherent decomposition rates using a reference temperature of 5 °C. Fast cycling C accounted for less than 5% of all C in both organic and mineral soils whereas the pool size of slow cycling C increased with C : N. Turnover time at 5 °C of fast cycling C typically was below 1 year, between 5 and 15 years for slow turning over C, and more than 500 years for passive C. We project that between 20 and 90% of the organic C could potentially be mineralized to CO2 within 50 incubation years at a constant temperature of 5 °C, with vulnerability to loss increasing in soils with higher C : N. These results demonstrate the variation in the vulnerability of C stored in permafrost soils based on inherent differences in organic matter decomposability, and point toward C : N as an index of decomposability that has the potential to be used to scale permafrost C loss across landscapes.
Schwartz E, Van Horn DJ, Buelow HN, Okie JG, Gooseff MN, Barrett JE, Takacs-Vesbach, CD (2014) Characterization of growing bacterial populations in McMurdo Dry Valley soils through stable isotope probing with 18O-water. FEMS Microbiology Letters 89(2): 415-425.
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Read PublicationSoil microbial communities of the McMurdo Dry Valleys, Antarctica (MDV) contain representatives from at least fourteen bacterial phyla. However, given low rates of microbial activity, it is unclear whether this richness represents functioning rather than dormant members of the community. We used stable isotope probing (SIP) with 18O-water to determine if microbial populations grow in MDV soils. Changes in the microbial community were characterized in soils amended with H2 18O and H2 18O-organic matter. Sequencing the 16S rRNA genes of the heavy and light fractions of the bacterial community DNA shows that DNA of microbial populations was labeled with 18O-water, indicating these micro-organisms grew in the MDV soils. Significant differences existed in the community composition of the heavy and light fractions of the H2 18O and H2 18O-organic matter amended samples (Anosim P < 0.05 of weighted Unifrac distance). Control samples and the light DNA fraction of the H2 18O amended samples were dominated by representatives of the phyla Deinococcus-Thermus, Proteobacteria, Planctomyces, Gemmatimonadetes, Actinobacteria and Acidobacteria, whereas Proteobacteria were more prevalent in the heavy DNA fractions from the H2 18O-water and the H2 18O-water-organic matter treatments. Our results indicate that SIP with H2 18O can be used to distinguish active bacterial populations even in this low organic matter environment.
Vogel JG, Bronson D, Gower ST, Schuur EAG (2014) The response of root and microbial respiration to the experimental warming of a boreal black spruce forest. Canadian Journal of Forest Research-Revue Canadienne De Recherche Forestiere 44(8): 986-993.
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Read PublicationWe investigated the effects of a 5 °C soil + air experimental heating on root and microbial respiration in a boreal black spruce (<i>Picea mariana</i> (Mill.) B.S.P.) forest in northern Manitoba, Canada, that was warmed between 2004 and 2007. In 2007, the <sup>14</sup>C/<sup>12</sup>C signatures of soil CO<sub>2</sub>efflux and root and soil microbial respiration were used in a two-pool mixing model to estimate their proportional contributions to soil CO<sub>2</sub> efflux and to examine how each changed in response to the warming treatments. In laboratory incubations, we examined whether warming had altered microbial respiration rates or microbial temperature sensitivity. The <sup>14</sup>C/<sup>12</sup>C signature of soil CO<sub>2</sub>efflux and microbial respiration in the heating treatments were both significantly (<i>p</i> < 0.05) enriched relative to the control treatment, suggesting that C deposited nearer the atmospheric bomb peak in 1963 contributed more to microbial respiration in heated than control treatments. Soil CO<sub>2</sub> efflux was significantly greater in the heated than control treatments, suggesting the acclimation to temperature of either root or microbial respiration was not occurring in 2007. Microbial respiration in laboratory incubations was similar in heated and control soils. This study shows that microbial respiration rates still responded to temperature even after 4 years of warming, highlighting that ecosystem warming can cause a prolonged release of soil organic matter from these soils.
Wojtowicz T, Compson ZG, Lamit LJ, Whitham TG and Gehring CA (2014) Plant genetic identity of foundation tree species and their hybrids affects a litter‑dwelling generalist predator. Oecologia 176(3): 799–810.
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Read PublicationThe effects of plant genetics on predators, especially those not living on the plant itself, are rarely studied and poorly understood. Therefore, we investigated the effect of plant hybridization and genotype on litter-dwelling spiders. Using an 18-year-old cottonwood common garden, we recorded agelenid sheet-web density associated with the litter layers of replicated genotypes of three tree cross types: Populus fremontii, Populus angustifolia, and their F1 hybrids. We surveyed 118 trees for agelenid litter webs at two distances from the trees (0–100 and 100–200 cm from trunk) and measured litter depth as a potential mechanism of web density patterns. Five major results emerged: web density within a 1-m radius of P. angustifolia was approximately three times higher than within a 1-m radius of P. fremontii, with F1 hybrids having intermediate densities; web density responded to P. angustifolia and F1 hybrid genotypes as indicated by a significant genotype × distance interaction, with some genotypes exhibiting a strong decline in web density with distance, while others did not; P. angustifolia litter layers were deeper than those of P. fremontii at both distance classes, and litter depth among P. angustifolia genotypes differed up to 300 %; cross type and genotype influenced web density via their effects on litter depth, and these effects were influenced by distance; web density was more sensitive to the effects of tree cross type than genotype. By influencing generalist predators, plant hybridization and genotype may indirectly impact trophic interactions such as intraguild predation, possibly affecting trophic cascades and ecosystem processes.
2013
Belshe EF, Schuur EAG, Bolker BM (2013) Tundra ecosystems observed to be CO2 sources due to differential amplification of the carbon cycle. Ecology Letters 16(10): 1307–1315.
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Read PublicationAre tundra ecosystems currently a carbon source or sink? What is the future trajectory of tundra carbon fluxes in response to climate change? These questions are of global importance because of the vast quantities of organic carbon stored in permafrost soils. In this meta-analysis, we compile 40 years of CO<sub>2</sub> flux observations from 54 studies spanning 32 sites across northern high latitudes. Using time-series analysis, we investigated if seasonal or annual CO<sub>2</sub> fluxes have changed over time, and whether spatial differences in mean annual temperature could help explain temporal changes in CO<sub>2</sub> flux. Growing season net CO<sub>2</sub> uptake has definitely increased since the 1990s; the data also suggest (albeit less definitively) an increase in winter CO<sub>2</sub> emissions, especially in the last decade. In spite of the uncertainty in the winter trend, we estimate that tundra sites were annual CO<sub>2</sub> sources from the mid-1980s until the 2000s, and data from the last 7 years show that tundra continue to emit CO<sub>2</sub> annually. CO<sub>2</sub> emissions exceed CO<sub>2</sub>uptake across the range of temperatures that occur in the tundra biome. Taken together, these data suggest that despite increases in growing season uptake, tundra ecosystems are currently CO<sub>2</sub> sources on an annual basis.
Belshe EF, Schuur EAG, Grosse G (2013) Quantification of upland thermokarst features with high resolution remote sensing. Environmental Research Letters 8(3): 035016.
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Read PublicationClimate-induced changes to permafrost are altering high latitude landscapes in ways that could increase the vulnerability of the vast soil carbon pools of the region. Permafrost thaw is temporally dynamic and spatially heterogeneous because, in addition to the thickening of the active layer, localized thermokarst features form when ice-rich permafrost thaws and the ground subsides. Thermokarst produces a diversity of landforms and alters the physical environment in dynamic ways. To estimate potential changes to the carbon cycle it is imperative to quantify the size and distribution of thermokarst landforms. By performing a supervised classification on a high resolution IKONOS image, we detected and mapped small, irregular thermokarst features occurring within an upland watershed in discontinuous permafrost of Interior Alaska. We found that 12% of the Eight Mile Lake (EML) watershed has undergone thermokarst, predominantly in valleys where tussock tundra resides. About 35% of the 3.7 km<sup>2</sup> tussock tundra class has likely transitioned to thermokarst. These landscape level changes created by permafrost thaw at EML have important implications for ecosystem carbon cycling because thermokarst features are forming in carbon-rich areas and are altering the hydrology in ways that increase seasonal thawing of the soil.
Compson ZG, KJ Adams, JA Edwards, JA Maestas, TG Whitham, and JC Marks (2013) Leaf litter quality affects aquatic insect emergence: contrasting patterns from two foundation trees. Oecologia 173 173(2): 507-519.
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Read PublicationReciprocal subsidies between rivers and terrestrial habitats are common where terrestrial leaf litter provides energy to aquatic invertebrates while emerging aquatic insects provide energy to terrestrial predators (e.g., birds, lizards, spiders). We examined how aquatic insect emergence changed seasonally with litter from two foundation riparian trees, whose litter often dominates riparian streams of the southwestern United States: Fremont (Populus fremontii) and narrowleaf (Populus angustifolia) cottonwood. P. fremontii litter is fast-decomposing and lower in defensive phytochemicals (i.e., condensed tannins, lignin) relative to P. angustifolia. We experimentally manipulated leaf litter from these two species by placing them in leaf enclosures with emergence traps attached in order to determine how leaf type influenced insect emergence. Contrary to our initial predictions, we found that packs with slow-decomposing leaves tended to support more emergent insects relative to packs with fast-decomposing leaves. Three findings emerged. Firstly, abundance (number of emerging insects m−2 day−1) was 25 % higher on narrowleaf compared to Fremont leaves for the spring but did not differ in the fall, demonstrating that leaf quality from two dominant trees of the same genus yielded different emergence patterns and that these patterns changed seasonally. Secondly, functional feeding groups of emerging insects differed between treatments and seasons. Specifically, in the spring collector-gatherer abundance and biomass were higher on narrowleaf leaves, whereas collector-filterer abundance and biomass were higher on Fremont leaves. Shredder abundance and biomass were higher on narrowleaf leaves in the fall. Thirdly, diversity (Shannon’s H′) was higher on Fremont leaves in the spring, but no differences were found in the fall, showing that fast-decomposing leaves can support a more diverse, complex emergent insect assemblage during certain times of the year. Collectively, these results challenge the notion that leaf quality is a simple function of decomposition, suggesting instead that aquatic insects benefit differentially from different leaf types, such that some use slow-decomposing litter for habitat and its temporal longevity and others utilize fast-decomposing litter with more immediate nutrient release.
Day FP, Schroeder RE, Stover DB, Brown ALP, Butnor JR, Dilustro J, Hungate BA, Dijkstra P, Duval BD, Seiler TJ, Drake BG, Hinkle CR (2013) The effects of 11 yr of CO2 enrichment on roots in a Florida scrub-oak ecosystem. New Phytologist 200(3): 778-787.
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Read Publication<ul id="nph12246-list-0001" class="u-list--bullet o-list--paragraph">
<li>Uncertainty surrounds belowground plant responses to rising atmospheric CO<sub>2</sub> because roots are difficult to measure, requiring frequent monitoring as a result of fine root dynamics and long-term monitoring as a result of sensitivity to resource availability.</li>
<li>We report belowground plant responses of a scrub-oak ecosystem in Florida exposed to 11 yr of elevated atmospheric CO<sub>2</sub> using open-top chambers. We measured fine root production, turnover and biomass using minirhizotrons, coarse root biomass using ground-penetrating radar and total root biomass using soil cores.</li>
<li>Total root biomass was greater in elevated than in ambient plots, and the absolute difference was larger than the difference aboveground. Fine root biomass fluctuated by more than a factor of two, with no unidirectional temporal trend, whereas leaf biomass accumulated monotonically. Strong increases in fine root biomass with elevated CO<sub>2</sub> occurred after fire and hurricane disturbance. Leaf biomass also exhibited stronger responses following hurricanes.</li>
<li>Responses after fire and hurricanes suggest that disturbance promotes the growth responses of plants to elevated CO<sub>2</sub>. Increased resource availability associated with disturbance (nutrients, water, space) may facilitate greater responses of roots to elevated CO<sub>2</sub>. The disappearance of responses in fine roots suggests limits on the capacity of root systems to respond to CO<sub>2</sub> enrichment.</li>
</ul>
Duval BD, Dijkstra P, Drake BG, Johnson DW, Ketterer ME, Megonigal JP, Hungate BA (2013) Element Pool Changes within a Scrub-Oak Ecosystem after 11 Years of Exposure to Elevated CO 2. PLOS ONE 8(5): e64386.
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Read PublicationThe effects of elevated CO<sub>2</sub> on ecosystem element stocks are equivocal, in part because cumulative effects of CO<sub>2</sub> on element pools are difficult to detect. We conducted a complete above and belowground inventory of non-nitrogen macro- and micronutrient stocks in a subtropical woodland exposed to twice-ambient CO<sub>2</sub> concentrations for 11 years. We analyzed a suite of nutrient elements and metals important for nutrient cycling in soils to a depth of ∼2 m, in leaves and stems of the dominant oaks, in fine and coarse roots, and in litter. In conjunction with large biomass stimulation, elevated CO<sub>2</sub> increased oak stem stocks of Na, Mg, P, K, V, Zn and Mo, and the aboveground pool of K and S. Elevated CO<sub>2</sub> increased root pools of most elements, except Zn. CO<sub>2</sub>-stimulation of plant Ca was larger than the decline in the extractable Ca pool in soils, whereas for other elements, increased plant uptake matched the decline in the extractable pool in soil. We conclude that elevated CO<sub>2</sub> caused a net transfer of a subset of nutrients from soil to plants, suggesting that ecosystems with a positive plant growth response under high CO<sub>2</sub> will likely cause mobilization of elements from soil pools to plant biomass.
Elberling B, Michelsen A, Schädel C, Schuur EAG, Christiansen HH, Berg L, Tamstorf MP, and Sigsgaard C (2013) Long-term CO2 production following permafrost thaw. Nature Climate Change 3(10): 890–894.
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Read PublicationThawing permafrost represents a poorly understood feedback mechanism of climate change in the Arctic, but with a potential impact owing to stored carbon being mobilized<sup><a id="ref-link-1" title="Tarnocai, C. et al. Soil organic carbon pools in the northern circumpolar permafrost region. Glob. Biogeochem. Cycle 23, GB2023 (2009)." href="http://www.nature.com/nclimate/journal/v3/n10/full/nclimate1955.html#ref1">1</a>, <a id="ref-link-2" title="Schuur, E. A. G. et al. The effect of permafrost thaw on old carbon release and net carbon exchange from tundra. Nature 459, 556-559 (2009)." href="http://www.nature.com/nclimate/journal/v3/n10/full/nclimate1955.html#ref2">2</a>, <a id="ref-link-3" title="Schuur, E. A. G. et al. Vulnerability of permafrost carbon to climate change: Implications for the global carbon cycle. BioScience 58, 701-714 (2008)." href="http://www.nature.com/nclimate/journal/v3/n10/full/nclimate1955.html#ref3">3</a>, <a id="ref-link-4" title="Hollesen, J., Elberling, B. & Jansson, P. E. Future active layer dynamics and CO2 production from thawing permafrost layers in Northeast Greenland. Glob. Change Biol. 17, 911-926 (2011)." href="http://www.nature.com/nclimate/journal/v3/n10/full/nclimate1955.html#ref4">4</a>, <a id="ref-link-5" title="Elberling, B., Christiansen, H. H. & Hansen, B. U. High nitrous oxide production from thawing permafrost. Nature Geosci. 3, 332-335 (2010)." href="http://www.nature.com/nclimate/journal/v3/n10/full/nclimate1955.html#ref5">5</a></sup>. We have quantified the long-term loss of carbon (C) from thawing permafrost in Northeast Greenland from 1996 to 2008 by combining repeated sediment sampling to assess changes in C stock and >12 years of CO<sub>2</sub> production in incubated permafrost samples. Field observations show that the active-layer thickness has increased by >1<span class="mb"><span class="mb"> </span></span>cm<span class="mb"><span class="mb"> </span></span>yr<sup>−1</sup> but thawing has not resulted in a detectable decline in C stocks. Laboratory mineralization rates at 5<span class="mb"><span class="mb"> </span></span>°C resulted in a C loss between 9 and 75<span class="mb">%</span>, depending on drainage, highlighting the potential of fast mobilization of permafrost C under aerobic conditions, but also that C at near-saturated conditions may remain largely immobilized over decades. This is confirmed by a three-pool C dynamics model that projects a potential C loss between 13 and 77<span class="mb">%</span> for 50 years of incubation at 5<span class="mb"><span class="mb"> </span></span>°C.
Hedlund BP, Paraiso JJ, Williams AJ, Huang Q, Wei Y, Dijkstra P, Hungate BA, Dong H, Zhang CL (2013) Wide distribution of autochthonous branched glycerol dialkyl glycerol tetraethers (bGDGTs) in US Great Basin hot springs. Frontiers in Microbiology 4.
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Read Publication<div data-canvas-width="78.4225">Branched glycerol dialkyl glycerol tetraethers (bGDGTs) are membrane-spanning lipids that likely stabilize membranes of some bacteria. Although bGDGTs have been reported previously in certain geothermal environments, it has been suggested that they may derive from surrounding soils since bGDGTs are known to be produced by soil bacteria. To test the hypothesis that bGDGTs can be produced by thermophiles in geothermal environments, we examined the distribution and abundance of bGDGTs, along with extensive geochemical data, in 40 sediment and mat samples collected from geothermal systems in the U.S. Great Basin (temperature: 31–95◦C; pH: 6.8–10.7). bGDGTs were found in 38 out of 40 samples at concentrations up to 824 ng/g sample dry mass and comprised up to 99.5% of total GDGTs (branched plus isoprenoidal). The wide distribution of bGDGTs in hot springs, strong correlation between core and polar lipid abundances, distinctness of bGDGT profiles compared to nearby soils, and higher concentration of bGDGTs in hot springs compared to nearby soils provided evidence of in situ production,</div>
<div data-canvas-width="19.563666666666666">particularly for the minimally methylated bGDGTs I, Ib, and Ic. Polar bGDGTs were found almost exclusively in samples ≤70◦C and the absolute abundance of polar bGDGTs correlated negatively with properties of chemically reduced, high temperature spring sources (temperature, H2S/HS−) and positively with properties of oxygenated, low temperature sites (O2, NO−3). Two-way cluster analysis and nonmetric multidimensional scaling based on relative abundance of polar bGDGTs supported these relationships and showed a negative relationship between the degree of methylation and temperature, suggesting a higher abundance for minimally methylated bGDGTs at high temperature. This study presents evidence of the widespread production of bGDGTs in mats and sediments of natural geothermal springs in the U.S. Great Basin, especially in oxygenated, low-temperature sites (70◦C).</div>
Hicks Pries CE, Schuur EAG, Crummer KG (2013) Thawing permafrost increases old soil and autotrophic respiration in tundra: Partitioning ecosystem respiration using δ13C and ∆14C. Global Change Biology 19(2):649-661.
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Read PublicationEcosystem respiration (<em>R</em><sub>eco</sub>) is one of the largest terrestrial carbon (<span class="fixed-roman">C</span>) fluxes. The effect of climate change on <em>R</em><sub>eco</sub> depends on the responses of its autotrophic and heterotrophic components. How autotrophic and heterotrophic respiration sources respond to climate change is especially important in ecosystems underlain by permafrost. Permafrost ecosystems contain vast stores of soil <span class="fixed-roman">C</span> (1672 Pg) and are located in northern latitudes where climate change is accelerated. Warming will cause a positive feedback to climate change if heterotrophic respiration increases without corresponding increases in primary production. We quantified the response of autotrophic and heterotrophic respiration to permafrost thaw across the 2008 and 2009 growing seasons. We partitioned <em>R</em><sub>eco</sub> using Δ<sup>14</sup><span class="fixed-roman">C</span> and δ<sup>13</sup><span class="fixed-roman">C</span> into four sources–two autotrophic (above – and belowground plant structures) and two heterotrophic (young and old soil). We sampled the Δ<sup>14</sup><span class="fixed-roman">C</span> and δ<sup>13</sup><span class="fixed-roman">C</span> of sources using incubations and the Δ<sup>14</sup><span class="fixed-roman">C</span> and δ<sup>13</sup><span class="fixed-roman">C</span> of <em>R</em><sub>eco</sub> using field measurements. We then used a Bayesian mixing model to solve for the most likely contributions of each source to <em>R</em><sub>eco</sub>. Autotrophic respiration ranged from 40 to 70% of <em>R</em><sub>eco</sub> and was greatest at the height of the growing season. Old soil heterotrophic respiration ranged from 6 to 18% of <em>R</em><sub>eco</sub> and was greatest where permafrost thaw was deepest. Overall, growing season fluxes of autotrophic and old soil heterotrophic respiration increased as permafrost thaw deepened. Areas with greater thaw also had the greatest primary production. Warming in permafrost ecosystems therefore leads to increased plant and old soil respiration that is initially compensated by increased net primary productivity. However, barring large shifts in plant community composition, future increases in old soil respiration will likely outpace productivity, resulting in a positive feedback to climate change.
Hicks Pries CE, Schuur EAG, Vogel JG, Natali SM (2013) Moisture drives surface decomposition in thawing tundra. Journal of Geophysical Research: Biogeosciences 118(3): 1133–1143.
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Read Publication<span class="paraNumber">[1]</span> Permafrost thaw can affect decomposition rates by changing environmental conditions and litter quality. As permafrost thaws, soils warm and thermokarst (ground subsidence) features form, causing some areas to become wetter while other areas become drier. We used a common substrate to measure how permafrost thaw affects decomposition rates in the surface soil in a natural permafrost thaw gradient and a warming experiment in Healy, Alaska. Permafrost thaw also changes plant community composition. We decomposed 12 plant litters in a common garden to test how changing plant litter inputs would affect decomposition. We combined species' tissue-specific decomposition rates with species and tissue-level estimates of aboveground net primary productivity to calculate community-weighted decomposition constants at both the thaw gradient and warming experiment. Moisture, specifically growing season precipitation and water table depth, was the most significant driver of decomposition. At the gradient, an increase in growing season precipitation from 200 to 300 mm increased mass loss of the common substrate by 100%. At the warming experiment, a decrease in the depth to the water table from 30 to 15 cm increased mass loss by 100%. At the gradient, community-weighted decomposition was 21% faster in extensive than in minimal thaw, but was similar when moss production was included. Overall, the effect of climate change and permafrost thaw on surface soil decomposition are driven more by precipitation and soil environment than by changes to plant communities. Increasing soil moisture is thereby another mechanism by which permafrost thaw can become a positive feedback to climate change.
Hou W, Wang S, Dong H, Jiang H, Briggs BR, Peacock JP, Huang Q, Huang L, Wu G, Zhi X, Li W, Dodsworth JA, Hedlund BP, Zhang C, Hartnett HE, Dijkstra P, Hungate BA (2013) A comprehensive census of microbial diversity in hot springs of Tengchong, Yunnan Province China using 16S rRNA gene pyrosequencing. PloS One 8(1): e53350.
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Read PublicationThe Rehai and Ruidian geothermal fields, located in Tengchong County, Yunnan Province, China, host a variety of geochemically distinct hot springs. In this study, we report a comprehensive, cultivation-independent census of microbial communities in 37 samples collected from these geothermal fields, encompassing sites ranging in temperature from 55.1 to 93.6°C, in pH from 2.5 to 9.4, and in mineralogy from silicates in Rehai to carbonates in Ruidian. Richness was low in all samples, with 21–123 species-level OTUs detected. The bacterial phylum <em>Aquificae</em> or archaeal phylum <em>Crenarchaeota</em> were dominant in Rehai samples, yet the dominant taxa within those phyla depended on temperature, pH, and geochemistry. Rehai springs with low pH (2.5–2.6), high temperature (85.1–89.1°C), and high sulfur contents favored the crenarchaeal order <em>Sulfolobales,</em> whereas those with low pH (2.6–4.8) and cooler temperature (55.1–64.5°C) favored the <em>Aquificae</em> genus <em>Hydrogenobaculum</em>. Rehai springs with neutral-alkaline pH (7.2–9.4) and high temperature (>80°C) with high concentrations of silica and salt ions (Na, K, and Cl) favored the <em>Aquificae</em> genus <em>Hydrogenobacter</em> and crenarchaeal orders <em>Desulfurococcales</em> and <em>Thermoproteales</em>. <em>Desulfurococcales</em> and <em>Thermoproteales</em> became predominant in springs with pH much higher than the optimum and even the maximum pH known for these orders. Ruidian water samples harbored a single <em>Aquificae</em> genus <em>Hydrogenobacter</em>, whereas microbial communities in Ruidian sediment samples were more diverse at the phylum level and distinctly different from those in Rehai and Ruidian water samples, with a higher abundance of uncultivated lineages, close relatives of the ammonia-oxidizing archaeon “<em>Candidatus</em> Nitrosocaldus yellowstonii”, and candidate division O1aA90 and OP1. These differences between Ruidian sediments and Rehai samples were likely caused by temperature, pH, and sediment mineralogy. The results of this study significantly expand the current understanding of the microbiology in Tengchong hot springs and provide a basis for comparison with other geothermal systems around the world.
Hungate BA, Day FP, Dijkstra P, Duval BD, Hinkle CR, Langley JA, Megonigal JP, Stiling P, Johnson DW, Drake BG (2013) Fire, hurricane and carbon dioxide: effects on net primary production of a subtropical woodland. New Phytologist 200(3): 767-777.
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Read Publication<ul id="nph12409-list-0001" class="u-list--bullet o-list--paragraph">
<li>Disturbance affects most terrestrial ecosystems and has the potential to shape their responses to chronic environmental change.</li>
<li>Scrub-oak vegetation regenerating from fire disturbance in subtropical Florida was exposed to experimentally elevated carbon dioxide (CO<sub>2</sub>) concentration (+350 μl l<sup>−1</sup>) using open-top chambers for 11 yr, punctuated by hurricane disturbance in year 8. Here, we report the effects of elevated CO<sub>2</sub> on aboveground and belowground net primary productivity (NPP) and nitrogen (N) cycling during this experiment.</li>
<li>The stimulation of NPP and N uptake by elevated CO<sub>2</sub> peaked within 2 yr after disturbance by fire and hurricane, when soil nutrient availability was high. The stimulation subsequently declined and disappeared, coincident with low soil nutrient availability and with a CO<sub>2</sub>-induced reduction in the N concentration of oak stems.</li>
<li>These findings show that strong growth responses to elevated CO<sub>2</sub> can be transient, are consistent with a progressively limited response to elevated CO<sub>2</sub> interrupted by disturbance, and illustrate the importance of biogeochemical responses to extreme events in modulating ecosystem responses to global environmental change.</li>
</ul>
Hungate BA, Dijkstra P, Wu Z, Duval BD, Day FP, Johnson DW, Megonigal JP, Brown ALP, Garland JL (2013) Cumulative response of ecosystem carbon and nitrogen stocks to chronic CO2 exposure in a subtropical oak woodland. New Phytologist 200(3): 753-766.
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Read Publication<ul id="nph12333-list-0001" class="u-list--bullet o-list--paragraph">
<li>Rising atmospheric carbon dioxide (CO<sub>2</sub>) could alter the carbon (C) and nitrogen (N) content of ecosystems, yet the magnitude of these effects are not well known. We examined C and N budgets of a subtropical woodland after 11 yr of exposure to elevated CO<sub>2</sub>.</li>
<li>We used open-top chambers to manipulate CO<sub>2</sub> during regrowth after fire, and measured C, N and tracer <sup>15</sup>N in ecosystem components throughout the experiment.</li>
<li>Elevated CO<sub>2</sub> increased plant C and tended to increase plant N but did not significantly increase whole-system C or N. Elevated CO<sub>2</sub> increased soil microbial activity and labile soil C, but more slowly cycling soil C pools tended to decline. Recovery of a long-term <sup>15</sup>N tracer indicated that CO<sub>2</sub> exposure increased N losses and altered N distribution, with no effect on N inputs.</li>
<li>Increased plant C accrual was accompanied by higher soil microbial activity and increased C losses from soil, yielding no statistically detectable effect of elevated CO<sub>2</sub> on net ecosystem C uptake. These findings challenge the treatment of terrestrial ecosystems responses to elevated CO<sub>2</sub> in current biogeochemical models, where the effect of elevated CO<sub>2</sub> on ecosystem C balance is described as enhanced photosynthesis and plant growth with decomposition as a first-order response.</li>
</ul>
Hurteau MD, Hungate BA, Koch GW, North MP, Smith GR (2013) Aligning ecology and markets in the forest carbon cycle. Frontiers in Ecology and the Environment 11(1): 37-42.
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Read PublicationA forest carbon (C) offset is a quantifiable unit of C that is commonly developed at the local or regional project scale and is designed to counterbalance anthropogenic C emissions by sequestering C in trees. In cap-and-trade programs, forest offsets have market value if the sequestered C is additional (more than would have occurred in the absence of the project) and permanent (sequestered within the project boundary for a specified period of time). Local management and ecological context determine the rate of C sequestration, risk of loss, and hence the market value. An understanding of global C dynamics can inform policy but may not be able to effectively price an ecosystem service, such as C sequestration. Appropriate pricing requires the assistance of ecologists to assess C stock abundance and stability over spatial and temporal scales appropriate for the regional market. We use the risk that sequestered C will be emitted as a result of wildfire (reversal risk) to show how ecological context can influence market valuation in offset programs.
Liu CM, Hungate BA, Tobian AAR, Serwadda D, Ravel J, Lester R, Kigozi G, Aziz M, Galiwango RM, Nalugoda F, Contente-Cuomo TL, Wawer MJ, Keim P, Gray RH, Price LB (2013) Male circumcision significantly reduces prevalence and load of genital anaerobic bacteria. MBio 4(2): e00076-13.
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Read PublicationMale circumcision reduces female-to-male HIV transmission. Hypothesized mechanisms for this protective effect include decreased HIV target cell recruitment and activation due to changes in the penis microbiome. We compared the coronal sulcus microbiota of men from a group of uncircumcised controls (<em>n</em> = 77) and from a circumcised intervention group (<em>n</em> = 79) at enrollment and year 1 follow-up in a randomized circumcision trial in Rakai, Uganda. We characterized microbiota using16S rRNA gene-based quantitative PCR (qPCR) and pyrosequencing, log response ratio (LRR), Bayesian classification, nonmetric multidimensional scaling (nMDS), and permutational multivariate analysis of variance (PerMANOVA). At baseline, men in both study arms had comparable coronal sulcus microbiota; however, by year 1, circumcision decreased the total bacterial load and reduced microbiota biodiversity. Specifically, the prevalence and absolute abundance of 12 anaerobic bacterial taxa decreased significantly in the circumcised men. While aerobic bacterial taxa also increased postcircumcision, these gains were minor. The reduction in anaerobes may partly account for the effects of circumcision on reduced HIV acquisition.
Millman JM, Waits K, Grande H, Marks AR, Marks JC, Price LB, and Hungate BA (2013) Prevalence of antibiotic-resistant E. coli in retail chicken: comparing conventional, organic, kosher, and raised without antibiotics. F1000 Research 2: 155.
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Read PublicationRetail poultry products are known sources of antibiotic-resistant <em>Escherichia coli</em>, a major human health concern. Consumers have a range of choices for poultry, including conventional, organic, kosher, and raised without antibiotics (RWA) – designations that are perceived to indicate differences in quality and safety. However, whether these categories vary in the frequency of contamination with antibiotic-resistant <em>E. coli</em> is unknown. We examined the occurrence of antibiotic-resistant <em>E. coli</em> on raw chicken marketed as conventional, organic, kosher and RWA. From April – June 2012, we purchased 213 samples of raw chicken from 15 locations in the New York City metropolitan area. We screened <em>E. coli</em> isolates from each sample for resistance to 12 common antibiotics. Although the organic and RWA labels restrict the use of antibiotics, the frequency of antibiotic-resistant <em>E. coli</em> tended to be only slightly lower for RWA, and organic chicken was statistically indistinguishable from conventional products that have no restrictions. Kosher chicken had the highest frequency of antibiotic-resistant <em>E. coli</em>, nearly twice that of conventional products, a result that belies the historical roots of kosher as a means to ensure food safety. These results indicate that production methods influence the frequency of antibiotic-resistant <em>E. coli </em>on poultry products available to consumers. Future research to identify the specific practices that cause the high frequency of antibiotic-resistant <em>E. coli</em> in kosher chicken could promote efforts to reduce consumer exposure to this potential pathogen.
Paraiso JJ, Williams AJ, Huang Q, Wei Y, Dijkstra P, Hungate BA, Dong H, Hedlund BP, Zhang CL (2013) The distribution and abundance of archaeal tetraether lipids in US Great Basin hot springs. Frontiers in Microbiology 4: 247.
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Read Publication<div data-canvas-width="203.71166666666667">Isoprenoidal glycerol dialkyl glycerol tetra ethers (iGDGTs) are core membrane lipids of many archaea that enhance the integrity of cytoplasmic membranes in extreme environments. We examined the iGDGT profiles and corresponding aqueous geochemistry in 40 hot spring sediment and microbial mat samples from the U.S. Great Basin with temperatures ranging from 31 to 95◦C and pH ranging from 6.8 to 10.7. The absolute abundance of iGDGTs correlated negatively with pH and positively with temperature. High lipid concentrations, distinct lipid profiles, and a strong relationship between polar and core lipids in hot spring samples suggested in situ production of most iGDGTs rather than contamination from local soils. Two-way cluster analysis and non-metric multidimensional scaling (NMS) of polar iGDGTs indicated that the relative abundance of individual lipids was most strongly related to temperature (r2=0.546), with moderate correlations with pH (r2=0.359), nitrite (r2=0.286), oxygen (r2=0.259), and nitrate(r2=0.215). Relative abundance profiles of individual polar iGDGTs indicated potential temperature optima for iGDGT-0 (≤70◦C), iGDGT-3 (≥55◦C), and iGDGT-4 (≥60◦C). These relationships likely reflect both physiological adaptations and community-level population shifts in response to temperature differences, such as a shift from cooler samples with more abundant methanogens to higher-temperature samples with more abundant Crenarchaeota. Crenarchaeol was widely distributed across the temperature gradient, which is consistent with other reports of abundant crenarchaeol in Great Basin hot springs and suggests a wide distribution for thermophilic ammonia-oxidizing archaea (AOA).</div>
Schädel C, Luo Y, Evans DR, Fei S, Schaeffer SM (2013) Separating soil CO2 efflux into C-pool-specific decay rates via inverse analysis of soil incubation data. Oecologia 171:721-732.
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Download .PDFSoil organic matter (SOM) is heterogeneous in structure and has been considered to consist of various pools with different intrinsic turnover rates. Although those pools have been conceptually expressed in models and analyzed according to soil physical and chemical properties, separation of SOM into component pools is still challenging. In this study, we conducted inverse analyses with data from a long-term (385 days) incubation experiment with two types of soil (from plant interspace and from underneath plants) to deconvolute soil carbon (C) efflux into different source pools. We analyzed the two datasets with one-, two- and three-pool models and used probability density functions as a criterion to judge the best model to fit the datasets. Our results indicated that soil C release trajectories over the 385 days of the incubation study were best modeled with a two-pool C model. For both soil types, released C within the first 10 days of the incubation study originated from the labile pool. Decomposition of C in the recalcitrant pool was modeled to contribute to the total CO<sub>2</sub> efflux by 9–11 % at the beginning of the incubation. At the end of the experiment, 75–85 % of the initial soil organic carbon (SOC) was modeled to be released over the incubation period. Our modeling analysis also indicated that the labile C-pool in the soil underneath plants was larger than that in soil from interspace. This deconvolution analysis was based on information contained in incubation data to separate carbon pools and can facilitate integration of results from incubation experiments into ecosystem models with improved parameterization.
Schuur EAG, Abbott BW, Bowden WB, Brovkin V, Camill P, Canadell JG, Chanton JP, Chapin III FS, Christensen TR, Ciais P, Crosby BT, Czimczik CI, Grosse G, Harden J, Hayes DJ, Hugelius G, Jastrow JD, Jones JB, Kleinen T, Koven CD, Krinner G, Kuhry P, Lawrence DM, McGuire AD, Natali SM, O’Donnell JA, Ping CL, Riley WJ, Rinke A, Romanovsky VE, Sannel ABK, Schädel C, Schaefer K, Sky J, Subin ZM, Tarnocai C, Turetsky MR, Waldrop MP, Walter Anthony KM, Wickland KP, Wilson CJ, Zimov SA (2013) Expert assessment of vulnerability of permafrost carbon to climate change. Climatic Change 119(2): 359-374.
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Read PublicationApproximately 1700 Pg of soil carbon (C) are stored in the northern circumpolar permafrost zone, more than twice as much C than in the atmosphere. The overall amount, rate, and form of C released to the atmosphere in a warmer world will influence the strength of the permafrost C feedback to climate change. We used a survey to quantify variability in the perception of the vulnerability of permafrost C to climate change. Experts were asked to provide quantitative estimates of permafrost change in response to four scenarios of warming. For the highest warming scenario (RCP 8.5), experts hypothesized that C release from permafrost zone soils could be 19–45 Pg C by 2040, 162–288 Pg C by 2100, and 381–616 Pg C by 2300 in CO<sub>2</sub> equivalent using 100-year CH<sub>4</sub> global warming potential (GWP). These values become 50 % larger using 20-year CH<sub>4</sub> GWP, with a third to a half of expected climate forcing coming from CH<sub>4</sub> even though CH<sub>4</sub> was only 2.3 % of the expected C release. Experts projected that two-thirds of this release could be avoided under the lowest warming scenario (RCP 2.6). These results highlight the potential risk from permafrost thaw and serve to frame a hypothesis about the magnitude of this feedback to climate change. However, the level of emissions proposed here are unlikely to overshadow the impact of fossil fuel burning, which will continue to be the main source of C emissions and climate forcing.
Stiling P, Moon D, Rossi A, Forkner R, Hungate BA, Day FP, Schroeder RE, Drake B (2013) Direct and legacy effects of long‐term elevated CO2 on fine root growth and plant–insect interactions. New Phytologist 200(3): 788-795.
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Read Publication<ul id="nph12295-list-0001" class="u-list--bullet o-list--paragraph">
<li>Increasing atmospheric CO<sub>2</sub> concentrations alter leaf physiology, with effects that cascade to communities and ecosystems. Yet, responses over cycles of disturbance and recovery are not well known, because most experiments span limited ecological time. We examined the effects of CO<sub>2</sub> on root growth, herbivory and arthropod biodiversity in a woodland from 1996 to 2006, and the legacy of CO<sub>2</sub> enrichment on these processes during the year after the CO<sub>2</sub> treatment ceased.</li>
<li>We used minirhizotrons to study root growth, leaf censuses to study herbivory and pitfall traps to determine the effects of elevated CO<sub>2</sub> on arthropod biodiversity.</li>
<li>Elevated CO<sub>2</sub> increased fine root biomass, but decreased foliar nitrogen and herbivory on all plant species. Insect biodiversity was unchanged in elevated CO<sub>2</sub>. Legacy effects of elevated CO<sub>2</sub> disappeared quickly as fine root growth, foliar nitrogen and herbivory levels recovered in the next growing season following the cessation of elevated CO<sub>2</sub>.</li>
<li>Although the effects of elevated CO<sub>2</sub> cascade through plants to herbivores, they do not reach other trophic levels, and biodiversity remains unchanged. The legacy of 10 yr of elevated CO<sub>2</sub> on plant–herbivore interactions in this system appear to be minimal, indicating that the effects of elevated CO<sub>2</sub> may not accumulate over cycles of disturbance and recovery.</li>
</ul>
Todd-Brown KEO, Randerson JT, Post WM, Hoffman FM, Tarnocai C, Schuur EAG, Allison SD (2013) Causes of variation in soil carbon simulations from CMIP5 Earth system models and comparison with observations. Biogeosciences 10(3): 1717-1736.
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Read PublicationStocks of soil organic carbon represent a large component of the carbon cycle that may participate in climate change feedbacks, particularly on decadal and centennial timescales. For Earth system models (ESMs), the ability to accurately represent the global distribution of existing soil carbon stocks is a prerequisite for accurately predicting future carbon–climate feedbacks. We compared soil carbon simulations from 11 model centers to empirical data from the Harmonized World Soil Database (HWSD) and the Northern Circumpolar Soil Carbon Database (NCSCD). Model estimates of global soil carbon stocks ranged from 510 to 3040 Pg C, compared to an estimate of 1260 Pg C (with a 95% confidence interval of 890–1660 Pg C) from the HWSD. Model simulations for the high northern latitudes fell between 60 and 820 Pg C, compared to 500 Pg C (with a 95% confidence interval of 380–620 Pg C) for the NCSCD and 290 Pg C for the HWSD. Global soil carbon varied 5.9 fold across models in response to a 2.6-fold variation in global net primary productivity (NPP) and a 3.6-fold variation in global soil carbon turnover times. Model–data agreement was moderate at the biome level (<i>R</i><sup>2</sup> values ranged from 0.38 to 0.97 with a mean of 0.75); however, the spatial distribution of soil carbon simulated by the ESMs at the 1° scale was not well correlated with the HWSD (Pearson correlation coefficients less than 0.4 and root mean square errors from 9.4 to 20.8 kg C m<sup>−2</sup>). In northern latitudes where the two data sets overlapped, agreement between the HWSD and the NCSCD was poor (Pearson correlation coefficient 0.33), indicating uncertainty in empirical estimates of soil carbon. We found that a reduced complexity model dependent on NPP and soil temperature explained much of the 1° spatial variation in soil carbon within most ESMs (<i>R</i><sup>2</sup> values between 0.62 and 0.93 for 9 of 11 model centers). However, the same reduced complexity model only explained 10% of the spatial variation in HWSD soil carbon when driven by observations of NPP and temperature, implying that other drivers or processes may be more important in explaining observed soil carbon distributions. The reduced complexity model also showed that differences in simulated soil carbon across ESMs were driven by differences in simulated NPP and the parameterization of soil heterotrophic respiration (inter-model <i>R</i><sup>2</sup> = 0.93), not by structural differences between the models. Overall, our results suggest that despite fair global-scale agreement with observational data and moderate agreement at the biome scale, most ESMs cannot reproduce grid-scale variation in soil carbon and may be missing key processes. Future work should focus on improving the simulation of driving variables for soil carbon stocks and modifying model structures to include additional processes.
Van Groenigen KJ, Forristal D, Jones M, Smyth N, Schwartz E, Hungate B, Dijkstra P (2013) Using metabolic tracer techniques to assess the impact of tillage and straw management on microbial carbon use efficiency in soil. Soil Biology and Biochemistry (66): 139-145.
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Read PublicationTillage practices and straw management can affect soil microbial activities with consequences for soil organic carbon (C) dynamics. Microorganisms metabolize soil organic C and in doing so gain energy and building blocks for biosynthesis, and release CO<sub>2</sub> to the atmosphere. Insight into the response of microbial metabolic processes and C use efficiency (CUE; microbial C produced per substrate C utilized) to management practices may therefore help to predict long term changes in soil C stocks. In this study, we assessed the effects of reduced (RT) and conventional tillage (CT) on the microbial central C metabolic network, using soil samples from a 12-year-old field experiment in an Irish winter wheat cropping system. Straw was removed from half of the RT and CT plots after harvest or incorporated into the soil in the other half, resulting in four treatment combinations. We added 1-<sup>13</sup>C and 2,3-<sup>13</sup>C pyruvate and 1-<sup>13</sup>C and U-<sup>13</sup>C glucose as metabolic tracer isotopomers to composite soil samples taken at two depths (0–15 cm and 15–30 cm) from each of the treatments and used the rate of position-specific respired <sup>13</sup>CO<sub>2</sub> to parameterize a metabolic model. Model outcomes were then used to calculate CUE of the microbial community. Whereas the composite samples differed in CUE, the changes were small, with values ranging between 0.757 and 0.783 across treatments and soil depth. Increases in CUE were associated with a reduced tricarboxylic acid cycle and reductive pentose phosphate pathway activity and increased consumption of metabolic intermediates for biosynthesis. Our results suggest that RT and straw incorporation do not substantially affect CUE.
Van Groenigen KJ, Van Kessel C, Hungate BA (2013) Increased greenhouse-gas intensity of rice production under future atmospheric conditions. Nature Climate Change 3(3): 288-291.
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Read PublicationIncreased atmospheric CO<sub>2</sub> and rising temperatures are expected to affect rice yields and greenhouse-gas (GHG) emissions from rice paddies<sup><a title="Van Groenigen, K. J., Osenberg, C. W. & Hungate, B. A. Increased soil emissions of potent greenhouse gases under increased atmospheric CO2. Nature 475, 214-216 (2011)." href="http://www.nature.com/nclimate/journal/v3/n3/full/nclimate1712.html#ref1">1</a>, <a title="Lobell, D. B. & Field, C. B. Global scale climate[mdash]crop yield relationships and the impacts of recent warming. Environ. Res. Lett. 2, 014002 (2007)." href="http://www.nature.com/nclimate/journal/v3/n3/full/nclimate1712.html#ref2">2</a>, <a title="Ainsworth, E. A. Rice production in a changing climate: A meta-analysis of responses to elevated carbon dioxide and elevated ozone concentration. Glob. Change Biol. 14, 1642-209 (2008)." href="http://www.nature.com/nclimate/journal/v3/n3/full/nclimate1712.html#ref3">3</a>, <a title="Peng, S. et al. Rice yields decline with higher night temperature from global warming. Proc. Natl Acad. Sci. USA 101, 9971-9975 (2004)." href="http://www.nature.com/nclimate/journal/v3/n3/full/nclimate1712.html#ref4">4</a></sup>. This is important, because rice cultivation is one of the largest human-induced sources of the potent GHG methane<sup><a title="EPA Global Anthropogenic non-CO2 Greenhouse Gas Emissions: 1990-2020, EPA 430-R-06-003 (United States Environmental Protection Agency, 2006)." href="http://www.nature.com/nclimate/journal/v3/n3/full/nclimate1712.html#ref5">5</a></sup> (CH<sub>4</sub>) and rice is the world’s second-most produced staple crop<sup><a title="http://faostat.fao.org/site/567/default.aspx[num]ancor." href="http://www.nature.com/nclimate/journal/v3/n3/full/nclimate1712.html#ref6">6</a></sup>. The need for meeting a growing global food demand<sup><a title="Cassman, K. G., Dobermann, A., Walters, D. T. & Yang, H. Meeting cereal demand while protecting natural resources and improving environmental quality. Annu. Rev. Environ. Resour. 28, 315-358 (2003)." href="http://www.nature.com/nclimate/journal/v3/n3/full/nclimate1712.html#ref7">7</a></sup> argues for assessing GHG emissions from croplands on the basis of yield rather than land area<sup><a title="Van Groenigen, J. W., Velthof, G. L., Oenema, O., van Groenigen, K. J. & van Kessel, C. Towards an agronomic assessment of N2O emissions: A case study for arable crops. Eur. J. Soil Sci. 61, 903-913 (2010)." href="http://www.nature.com/nclimate/journal/v3/n3/full/nclimate1712.html#ref8">8</a>, <a title="Mosier, A. R., Halvorson, A. D., Reule, C.A. & Liu, X. J. J. Net global warming potential and greenhouse gas intensity in irrigated cropping systems in northeastern Colorado. J. Environ. Qual. 35, 1584-1598 (2006)." href="http://www.nature.com/nclimate/journal/v3/n3/full/nclimate1712.html#ref9">9</a>, <a title="Grassini, P. & Cassman, K. G. High-yield maize with large net energy yield and small global warming intensity. Proc. Natl Acad. USA 109, 1074-1079 (2012)." href="http://www.nature.com/nclimate/journal/v3/n3/full/nclimate1712.html#ref10">10</a></sup>, such that efforts to reduce GHG emissions take into consideration the consequences for food production. However, it is unclear whether or how the GHG intensity (that is, yield-scaled GHG emissions) of cropping systems will be affected by future atmospheric conditions. Here we show, using meta-analysis, that increased atmospheric CO<sub>2</sub> (ranging from 550 to 743<span class="mb"><span class="mb"> </span></span>ppmV) and warming (ranging from +0.8<span class="mb"><span class="mb"> </span></span>°C to +6<span class="mb"><span class="mb"> </span></span>°C) both increase the GHG intensity of rice cultivation. Increased atmospheric CO<sub>2</sub> increased GHG intensity by 31.4<span class="mb">%</span>, because CH<sub>4</sub> emissions are stimulated more than rice yields. Warming increased GHG intensity by 11.8<span class="mb">%</span> per 1<span class="mb"><span class="mb"> </span></span>°C, largely owing to a decrease in yield. This analysis suggests that rising CO<sub>2</sub> and warming will approximately double the GHG intensity of rice production by the end of the twenty-first century, stressing the need for management practices that optimize rice production while reducing its GHG intensity as the climate continues to change.
2012
Belshe EF, Schuur EAG, Bolker BM, Bracho R (2012) Incorporating spatial heterogeneity created by permafrost thaw into a landscape carbon estimate. Journal of Geophysical Research-Biogeosciences 117(G1).
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Read PublicationThe future carbon balance of high-latitude ecosystems is dependent on the sensitivity of biological processes (photosynthesis and respiration) to the physical changes occurring with permafrost thaw. Predicting C exchange in these ecosystems is difficult because the thawing of permafrost is a heterogeneous process that creates a complex landscape. We measured net ecosystem exchange of C using eddy covariance (EC) in a tundra landscape visibly undergoing thaw during two 6 month campaigns in 2008 and 2009. We developed a spatially explicit quantitative metric of permafrost thaw based on variation in microtopography and incorporated it into an EC carbon flux estimate using a generalized additive model (GAM). This model allowed us to make predictions about C exchange for the landscape as a whole and for specific landscape patches throughout the continuum of permafrost thaw and ground subsidence. During June through November 2008, the GAM predicted that the landscape on average took up 337.1 g C m<sup>−2</sup> via photosynthesis and released 289.5 g C m<sup>−2</sup> via respiration, resulting in a net C gain of 47.5 g C m<sup>−2</sup> by the tundra ecosystem. During April through October 2009, the landscape on average took up 498.7 g C m<sup>−2</sup> and released 410.3 g C m<sup>−2</sup>, resulting in a net C gain of 87.8 g C m<sup>−2</sup>. On average, between the years, areas with the highest permafrost thaw and ground subsidence photosynthesized 17.7% more and respired 3.3% more C than the average landscape. Areas with the least thaw and subsidence photosynthesized 15% less and respired 5.1% less than the landscape on average. By incorporating spatial variation into the EC C estimate, we were able to estimate the C balance of a heterogeneous landscape and determine the collective effect of permafrost thaw on the plant and soil processes that drive ecosystem C flux. In these study years, permafrost thaw appeared to increase the amplitude of the C cycle by stimulating both C release and sequestration, while the ecosystem remained a C sink at the landscape scale.
Brown JR, Blankinship JC, Niboyet A, van Groenigen KJ, Dijkstra P, Le Roux X, Leadley PW, Hungate BA (2012) Effects of multiple global change treatments on soil N2O fluxes. Biogeochemistry 109(1): 85-100.
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Read PublicationGlobal environmental changes are expected to alter ecosystem carbon and nitrogen cycling, but the interactive effects of multiple simultaneous environmental changes are poorly understood. Effects of these changes on the production of nitrous oxide (N<sub>2</sub>O), an important greenhouse gas, could accelerate climate change. We assessed the responses of soil N<sub>2</sub>O fluxes to elevated CO<sub>2</sub>, heat, altered precipitation, and enhanced nitrogen deposition, as well as their interactions, in an annual grassland at the Jasper Ridge Global Change Experiment (CA, USA). Measurements were conducted after 6, 7 and 8 years of treatments. Elevated precipitation increased N<sub>2</sub>O efflux, especially in combination with added nitrogen and heat. Path analysis supported the idea that increased denitrification due to increased soil water content and higher labile carbon availability best explained increased N<sub>2</sub>O efflux, with a smaller, indirect contribution from nitrification. In our data and across the literature, single-factor responses tended to overestimate interactive responses, except when global change was combined with disturbance by fire, in which case interactive effects were large. Thus, for chronic global environmental changes, higher order interactions dampened responses of N<sub>2</sub>O efflux to multiple global environmental changes, but interactions were strongly positive when global change was combined with disturbance. Testing whether these responses are general should be a high priority for future research.
Clingenpeel SR, Moan JL, McGrath DM, Hungate BA, Watwood ME (2012) Stable carbon isotope fractionation in chlorinated ethene degradation by bacteria expressing three toluene oxygenases. Frontiers in microbiology 3: 63.
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Read PublicationOne difficulty in using bioremediation at a contaminated site is demonstrating that biodegradation is actually occurring <em>in situ</em>. The stable isotope composition of contaminants may help with this, since they can serve as an indicator of biological activity. To use this approach it is necessary to establish how a particular biodegradation pathway affects the isotopic composition of a contaminant. This study examined bacterial strains expressing three aerobic enzymes for their effect on the <sup>13</sup>C/<sup>12</sup>C ratio when degrading both trichloroethene (TCE) and <em>cis</em>-1,2-dichloroethene (c-DCE): toluene 3-monoxygenase, toluene 4-monooxygenase, and toluene 2,3-dioxygenase. We found no significant differences in fractionation among the three enzymes for either compound. Aerobic degradation of c-DCE occurred with low fractionation producing δ<sup>13</sup>C enrichment factors of −0.9 ± 0.5 to −1.2 ± 0.5, in contrast to reported anaerobic degradation δ<sup>13</sup>C enrichment factors of −14.1 to −20.4‰. Aerobic degradation of TCE resulted in δ<sup>13</sup>C enrichment factors of −11.6 ± 4.1 to −14.7 ± 3.0‰ which overlap reported δ<sup>13</sup>C enrichment factors for anaerobic TCE degradation of −2.5 to −13.8‰. The data from this study suggest that stable isotopes could serve as a diagnostic for detecting aerobic biodegradation of TCE by toluene oxygenases at contaminated sites.
Dore S, Montes‐Helu M, Hart SC, Hungate BA, Koch GW, Moon JB, Finkral AJ, Kolb TE (2012) Recovery of ponderosa pine ecosystem carbon and water fluxes from thinning and stand‐replacing fire. Global Change Biology 18(10): 3171-3185.
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Read PublicationCarbon uptake by forests is a major sink in the global carbon cycle, helping buffer the rising concentration of CO<sub>2</sub> in the atmosphere, yet the potential for future carbon uptake by forests is uncertain. Climate warming and drought can reduce forest carbon uptake by reducing photosynthesis, increasing respiration, and by increasing the frequency and intensity of wildfires, leading to large releases of stored carbon. Five years of eddy covariance measurements in a ponderosa pine (<em>Pinus ponderosa</em>)-dominated ecosystem in northern Arizona showed that an intense wildfire that converted forest into sparse grassland shifted site carbon balance from sink to source for at least 15 years after burning. In contrast, recovery of carbon sink strength after thinning, a management practice used to reduce the likelihood of intense wildfires, was rapid. Comparisons between an undisturbed-control site and an experimentally thinned site showed that thinning reduced carbon sink strength only for the first two posttreatment years. In the third and fourth posttreatment years, annual carbon sink strength of the thinned site was higher than the undisturbed site because thinning reduced aridity and drought limitation to carbon uptake. As a result, annual maximum gross primary production occurred when temperature was 3 °C higher at the thinned site compared with the undisturbed site. The severe fire consistently reduced annual evapotranspiration (range of 12–30%), whereas effects of thinning were smaller and transient, and could not be detected in the fourth year after thinning. Our results show large and persistent effects of intense fire and minor and short-lived effects of thinning on southwestern ponderosa pine ecosystem carbon and water exchanges.
Dunbar J, Eichorst SA, Gallegos‐Graves LV, Silva S, Xie G, Hengartner NW, Evans RD, Hungate BA, Jackson RB, Megonigal JP, Schadt CW, Vilgalys R, Zak DR, Kuske CR (2012) Common bacterial responses in six ecosystems exposed to 10 years of elevated atmospheric carbon dioxide. Environmental Microbiology 14(5): 1145-1158.
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Read PublicationSix terrestrial ecosystems in the USA were exposed to elevated atmospheric CO<sub>2</sub> in single or multifactorial experiments for more than a decade to assess potential impacts. We retrospectively assessed soil bacterial community responses in all six-field experiments and found ecosystem-specific and common patterns of soil bacterial community response to elevated CO<sub>2</sub>. Soil bacterial composition differed greatly across the six ecosystems. No common effect of elevated atmospheric CO<sub>2</sub> on bacterial biomass, richness and community composition across all of the ecosystems was identified, although significant responses were detected in individual ecosystems. The most striking common trend across the sites was a decrease of up to 3.5-fold in the relative abundance of <em>Acidobacteria</em> Group 1 bacteria in soils exposed to elevated CO<sub>2</sub> or other climate factors. The <em>Acidobacteria</em> Group 1 response observed in exploratory 16S rRNA gene clone library surveys was validated in one ecosystem by 100-fold deeper sequencing and semi-quantitative PCR assays. Collectively, the 16S rRNA gene sequencing approach revealed influences of elevated CO<sub>2</sub> on multiple ecosystems. Although few common trends across the ecosystems were detected in the small surveys, the trends may be harbingers of more substantive changes in less abundant, more sensitive taxa that can only be detected by deeper surveys.
Duval BD, Blankinship JC, Dijkstra P, Hungate BA (2012) RETRACTED ARTICLE: CO2 effects on plant nutrient concentration depend on plant functional group and available nitrogen: a meta-analysis. Plant Ecology 213(3): 505-521.
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Read PublicationElevated CO<sub>2</sub> is expected to lower plant nutrient concentrations via carbohydrate dilution and increased nutrient use efficiency. Elevated CO<sub>2</sub> consistently lowers plant foliar nitrogen, but there is no consensus on CO<sub>2</sub> effects across the range of plant nutrients. We used meta-analysis to quantify elevated CO<sub>2</sub> effects on leaf, stem, root, and seed concentrations of B, Ca, Cu, Fe, K, Mg, Mn, P, S, and Zn among four plant functional groups and two levels of N fertilization. CO<sub>2</sub> effects on plant nutrient concentration depended on the nutrient, plant group, tissue, and N status. CO<sub>2</sub> reduced B, Cu, Fe, and Mg, but increased Mn concentration in the leaves of N<sub>2</sub> fixers. Elevated CO<sub>2</sub> increased Cu, Fe, and Zn, but lowered Mn concentration in grass leaves. Tree leaf responses were strongly related to N status: CO<sub>2</sub> significantly decreased Cu, Fe, Mg, and S at high N, but only Fe at low N. Elevated CO<sub>2</sub> decreased Mg and Zn in crop leaves grown with high N, and Mn at low N. Nutrient concentrations in crop roots were not affected by CO<sub>2</sub> enrichment, but CO<sub>2</sub> decreased Ca, K, Mg and P in tree roots. Crop seeds had lower S under elevated CO<sub>2</sub>. We also tested the validity of a “dilution model.” CO<sub>2</sub> reduced the concentration of plant nutrients 6.6% across nutrients and plant groups, but the reduction is less than expected (18.4%) from carbohydrate accumulation alone. We found that elevated CO<sub>2</sub> impacts plant nutrient status differently among the nutrient elements, plant functional groups, and among plant tissues. Our synthesis suggests that differences between plant groups and plant organs, N status, and differences in nutrient chemistry in soils preclude a universal hypothesis strictly related to carbohydrate dilution regarding plant nutrient response to elevated CO<sub>2</sub>.
Fountain AG, Campbell JL, Schuur EAG, Stammerjohn SE, Williams MW, Ducklow HW (2012) The Disappearing Cryosphere: Impacts and Ecosystem Responses to Rapid Cryosphere Loss. Bioscience 62:405-415.
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Read PublicationThe cryosphere—the portion of the Earth’s surface where water is in solid form for at least one month of the year—has been shrinking in response to climate warming. The extents of sea ice, snow, and glaciers, for example, have been decreasing. In response, the ecosystems within the cryosphere and those that depend on the cryosphere have been changing. We identify two principal aspects of ecosystem-level responses to cryosphere loss: (1) trophodynamic alterations resulting from the loss of habitat and species loss or replacement and (2) changes in the rates and mechanisms of biogeochemical storage and cycling of carbon and nutrients, caused by changes in physical forcings or ecological community functioning. These changes affect biota in positive or negative ways, depending on how they interact with the cryosphere. The important outcome, however, is the change and the response the human social system (infrastructure, food, water, recreation) will have to that change.
Harden JW, Koven CD, Ping CL, Hugelius G, McGuire AD, Camill P, Jorgenson T, Kuhry P, Michaelson GJ, O'Donnell JA, Schuur EAG, Tarnocai C, Johnson K, Grosse G (2012) Field information links permafrost carbon to physical vulnerabilities of thawing. Geophysical Research Letters 39: L15704.
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Read PublicationDeep soil profiles containing permafrost (Gelisols) were characterized for organic carbon (C) and total nitrogen (N) stocks to 3 m depths. Using the Community Climate System Model (CCSM4) we calculate cumulative distributions of active layer thickness (ALT) under current and future climates. The difference in cumulative ALT distributions over time was multiplied by C and N contents of soil horizons in Gelisol suborders to calculate newly thawed C and N. Thawing ranged from 147 PgC with 10 PgN by 2050 (representative concentration pathway RCP scenario 4.5) to 436 PgC with 29 PgN by 2100 (RCP 8.5). Organic horizons that thaw are vulnerable to combustion, and all horizon types are vulnerable to shifts in hydrology and decomposition. The rates and extent of such losses are unknown and can be further constrained by linking field and modelling approaches. These changes have the potential for strong additional loading to our atmosphere, water resources, and ecosystems.
Hooper DU, Adair EC, Cardinale BJ, Byrnes JEK, Hungate BA, Matulich KL, Gonzalez A, Duffy JE, Gamfeldt L, O’Connor MI (2012) A global synthesis reveals biodiversity loss as a major driver of ecosystem change. Nature 486(7401): 105-108.
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Read PublicationEvidence is mounting that extinctions are altering key processes important to the productivity and sustainability of Earth’s ecosystems<sup><a id="ref-link-1" title="Loreau, M., Naeem, S. & Inchausti, P. Biodiversity and Ecosystem Functioning: Synthesis and perspectives (Oxford Univ. Press, 2002)" href="http://www.nature.com/nature/journal/v486/n7401/full/nature11118.html#ref1">1</a>, <a id="ref-link-2" title="Hooper, D. U. et al. Effects of biodiversity on ecosystem functioning: a consensus of current knowledge. Ecol. Monogr. 75, 3-35 (2005)" href="http://www.nature.com/nature/journal/v486/n7401/full/nature11118.html#ref2">2</a>, <a id="ref-link-3" title="Tilman, D. Ecological consequences of biodiversity: a search for general principles. Ecology 80, 1455-1474 (1999)" href="http://www.nature.com/nature/journal/v486/n7401/full/nature11118.html#ref3">3</a>, <a id="ref-link-4" title="Wardle, D. A., Bardgett, R. D., Callaway, R. M. & Van der Putten, W. H. Terrestrial ecosystem responses to species gains and losses. Science 332, 1273-1277 (2011)" href="http://www.nature.com/nature/journal/v486/n7401/full/nature11118.html#ref4">4</a></sup>. Further species loss will accelerate change in ecosystem processes<sup><a id="ref-link-5" title="Balvanera, P. et al. Quantifying the evidence for biodiversity effects on ecosystem functioning and services. Ecol. Lett. 9, 1146-1156 (2006)" href="http://www.nature.com/nature/journal/v486/n7401/full/nature11118.html#ref5">5</a>, <a id="ref-link-6" title="Cardinale, B. J. et al. The functional role of producer diversity in ecosystems. Am. J. Bot. 98, 572-592 (2011)" href="http://www.nature.com/nature/journal/v486/n7401/full/nature11118.html#ref6">6</a>, <a id="ref-link-7" title="Stachowicz, J. J., Bruno, J. F. & Duffy, J. E. Understanding the effects of marine biodiversity on communities and ecosystems. Annu. Rev. Ecol. Evol. Syst. 38, 739-766 (2007)" href="http://www.nature.com/nature/journal/v486/n7401/full/nature11118.html#ref7">7</a>, <a id="ref-link-8" title="Perrings, C. et al. Ecosystem services, targets, and indicators for the conservation and sustainable use of biodiversity. Front. Ecol. Environ 9, 512-520 (2011)" href="http://www.nature.com/nature/journal/v486/n7401/full/nature11118.html#ref8">8</a></sup>, but it is unclear how these effects compare to the direct effects of other forms of environmental change that are both driving diversity loss and altering ecosystem function. Here we use a suite of meta-analyses of published data to show that the effects of species loss on productivity and decomposition—two processes important in all ecosystems—are of comparable magnitude to the effects of many other global environmental changes. In experiments, intermediate levels of species loss (21–40%) reduced plant production by 5–10%, comparable to previously documented effects of ultraviolet radiation and climate warming. Higher levels of extinction (41–60%) had effects rivalling those of ozone, acidification, elevated CO<sub>2</sub> and nutrient pollution. At intermediate levels, species loss generally had equal or greater effects on decomposition than did elevated CO<sub>2</sub> and nitrogen addition. The identity of species lost also had a large effect on changes in productivity and decomposition, generating a wide range of plausible outcomes for extinction. Despite the need for more studies on interactive effects of diversity loss and environmental changes, our analyses clearly show that the ecosystem consequences of local species loss are as quantitatively significant as the direct effects of several global change stressors that have mobilized major international concern and remediation efforts<sup><a id="ref-link-9" title="IPCC. Climate Change 2007: Synthesis Report. Contribution of Working Groups I, II and III to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change (eds Core Writing Team, Pachauri, R. K. & Reisinger, A.) (IPCC, 2007)" href="http://www.nature.com/nature/journal/v486/n7401/full/nature11118.html#ref9">9</a></sup>.
Lee H, Schuur EAG, Inglett KS, Lavoie M, Chanton JP (2012) The rate of permafrost carbon release under aerobic and anaerobic conditions and its potential effects on climate. Global Change Biology 18:515-527.
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Read PublicationRecent observations suggest that permafrost thaw may create two completely different soil environments: aerobic in relatively well-drained uplands and anaerobic in poorly drained wetlands. The soil oxygen availability will dictate the rate of permafrost carbon release as carbon dioxide (<span class="fixed-roman">CO</span><sub>2</sub>) and as methane (<span class="fixed-roman">CH</span><sub>4</sub>), and the overall effects of these emitted greenhouse gases on climate. The objective of this study was to quantify <span class="fixed-roman">CO</span><sub>2</sub> and <span class="fixed-roman">CH</span><sub>4</sub> release over a 500-day period from permafrost soil under aerobic and anaerobic conditions in the laboratory and to compare the potential effects of these emissions on future climate by estimating their relative climate forcing. We used permafrost soils collected from Alaska and Siberia with varying organic matter characteristics and simultaneously incubated them under aerobic and anaerobic conditions to determine rates of <span class="fixed-roman">CO</span><sub>2</sub> and <span class="fixed-roman">CH</span><sub>4</sub> production. Over 500 days of soil incubation at 15 °C, we observed that carbon released under aerobic conditions was 3.9–10.0 times greater than anaerobic conditions. When scaled by greenhouse warming potential to account for differences between <span class="fixed-roman">CO</span><sub>2</sub> and <span class="fixed-roman">CH</span><sub>4</sub>, relative climate forcing ranged between 1.5 and 7.1. Carbon release in organic soils was nearly 20 times greater than mineral soils on a per gram soil basis, but when compared on a per gram carbon basis, deep permafrost mineral soils showed carbon release rates similar to organic soils for some soil types. This suggests that permafrost carbon may be very labile, but that there are significant differences across soil types depending on the processes that controlled initial permafrost carbon accumulation within a particular landscape. Overall, our study showed that, independent of soil type, permafrost carbon in a relatively aerobic upland ecosystems may have a greater effect on climate when compared with a similar amount of permafrost carbon thawing in an anaerobic environment, despite the release of <span class="fixed-roman">CH</span><sub>4</sub> that occurs in anaerobic conditions.
Natali SM, Schuur EAG, Rubin RL (2012) Increased plant productivity in Alaskan tundra as a result of experimental warming of soil and permafrost. Journal of Ecology 100:488-498.
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Read PublicationThe response of northern tundra plant communities to warming temperatures is of critical concern because permafrost ecosystems play a key role in global carbon (C) storage, and climate-induced ecological shifts in the plant community will affect the transfer of carbon-dioxide between biological and atmospheric pools.
This study, which focuses on the response of tundra plant growth and phenology to experimental warming, was conducted at the Carbon in Permafrost Experimental Heating Research project, located in the northern foothills of the Alaska Range. We used snow fences coupled with spring snow removal to increase deep-soil temperatures and thaw depth (winter warming), and open-top chambers to increase summer air temperatures (summer warming).
Winter warming increased wintertime soil temperature (5–40 cm) by 2.3 °C, resulting in a 10% increase in growing season thaw depth. Summer warming significantly increased growing season air temperature; peak temperature differences occurred near midday when summer warming plots were approximately 1.0 °C warmer than ambient plots.
Changes in the soil environment as a result of winter warming treatment resulted in a 20% increase in above-ground biomass and net primary productivity (ANPP), while there was no detected summer warming effect on ecosystem-level ANPP or biomass. Both summer and winter warming extended the growing season through earlier bud break and delayed senescence, despite equivalent snow-free days across treatments. As with ANPP, winter warming increased canopy N mass by 20%, while there was no summer warming effect on canopy N.
The warming-mediated increase in N availability, coupled with phenological shifts, may have driven higher rates of ANPP in the winter warming plots, and the lack of ecosystem-level N and ANPP response to summer warming suggest continued N limitation in the summer warming plots.
Synthesis: These results highlight the role of soil and permafrost dynamics in regulating plant response to climate change and provide evidence that warming may promote greater C accumulation in tundra plant biomass. While warming temperatures are expected to enhance microbial decomposition of the large pool of organic matter stored in tundra soils and permafrost, these respiratory losses may be offset, at least in part, by warming-mediated increases in plant growth.
Pries CEH, Schuur EAG, Crummer KG (2012) Holocene Carbon Stocks and Carbon Accumulation Rates Altered in Soils Undergoing Permafrost Thaw. Ecosystems 15:162-173.
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Read PublicationPermafrost soils are a significant global store of carbon (C) with the potential to become a large C source to the atmosphere. Climate change is causing permafrost to thaw, which can affect primary production and decomposition, therefore affecting ecosystem C balance. To understand future responses of permafrost soils to climate change, we inventoried current soil C stocks, investigated ∆<sup>14</sup>C, C:N, δ<sup>13</sup>C, and δ<sup>15</sup>N depth profiles, modeled soil C accumulation rates, and calculated decadal net ecosystem production (NEP) in subarctic tundra soils undergoing minimal, moderate, and extensive permafrost thaw near Eight Mile Lake (EML) in Healy, Alaska. We modeled decadal and millennial soil C inputs, decomposition constants, and C accumulation rates by plotting cumulative C inventories against C ages based on radiocarbon dating of surface and deep soils, respectively. Soil C stocks at EML were substantial, over 50 kg C m<sup>−2</sup> in the top meter, and did not differ much among sites. Carbon to nitrogen ratio, δ<sup>13</sup>C, and δ<sup>15</sup>N depth profiles indicated most of the decomposition occurred within the organic soil horizon and practically ceased in deeper, frozen horizons. The average C accumulation rate for EML surface soils was 25.8 g C m<sup>−2</sup> y<sup>−1</sup> and the rate for the deep soil accumulation was 2.3 g C m<sup>−2</sup> y<sup>−1</sup>, indicating these systems have been C sinks throughout the Holocene. Decadal net ecosystem production averaged 14.4 g C m<sup>−2</sup> y<sup>−1</sup>. However, the shape of decadal C accumulation curves, combined with recent annual NEP measurements, indicates soil C accumulation has halted and the ecosystem may be becoming a C source. Thus, the net impact of climate warming on tundra ecosystem C balance includes not only becoming a C source but also the loss of C uptake capacity these systems have provided over the past ten thousand years.
Templer PH, Mack MC, Chapin III FS, Christenson LM, Compton JE, Crook HD, Currie WS, Curtis CJ, Dail DB, D'Antonio CM, Emmett BA, Epstein HE, Goodale CL, Gundersen P, Hobbie SE, Holland K, Hooper DU, Hungate BA, Lamontagne S, Nadelhoffer KJ, Osenberg CW, Perakis SS, Schleppi P, Schimel J, Schmidt IK, Sommerkorn M, Spoelstra J, Tietema A, Wessel WW, Zak DR (2012) Sinks for nitrogen inputs in terrestrial ecosystems: a meta-analysis of 15N tracer field studies. Ecology 93(8): 1816-1829.
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Read PublicationEffects of anthropogenic nitrogen (N) deposition and the ability of terrestrial ecosystems to store carbon (C) depend in part on the amount of N retained in the system and its partitioning among plant and soil pools. We conducted a meta-analysis of studies at 48 sites across four continents that used enriched <sup>15</sup>N isotope tracers in order to synthesize information about total ecosystem N retention (i.e., total ecosystem <sup>15</sup>N recovery in plant and soil pools) across natural systems and N partitioning among ecosystem pools. The greatest recoveries of ecosystem <sup>15</sup>N tracer occurred in shrublands (mean, 89.5%) and wetlands (84.8%) followed by forests (74.9%) and grasslands (51.8%). In the short term (<1 week after <sup>15</sup>N tracer application), total ecosystem <sup>15</sup>N recovery was negatively correlated with fine-root and soil <sup>15</sup>N natural abundance, and organic soil C and N concentration but was positively correlated with mean annual temperature and mineral soil C:N. In the longer term (3–18 months after <sup>15</sup>N tracer application), total ecosystem <sup>15</sup>N retention was negatively correlated with foliar natural-abundance <sup>15</sup>N but was positively correlated with mineral soil C and N concentration and C : N, showing that plant and soil natural-abundance <sup>15</sup>N and soil C:N are good indicators of total ecosystem N retention. Foliar N concentration was not significantly related to ecosystem <sup>15</sup>N tracer recovery, suggesting that plant N status is not a good predictor of total ecosystem N retention. Because the largest ecosystem sinks for <sup>15</sup>N tracer were below ground in forests, shrublands, and grasslands, we conclude that growth enhancement and potential for increased C storage in aboveground biomass from atmospheric N deposition is likely to be modest in these ecosystems. Total ecosystem <sup>15</sup>N recovery decreased with N fertilization, with an apparent threshold fertilization rate of 46 kg N·ha<sup>−1</sup>·yr<sup>−1</sup> above which most ecosystems showed net losses of applied <sup>15</sup>N tracer in response to N fertilizer addition.
Trucco C, Schuur EAG, Natali SM, Belshe EF, Bracho R, Vogel J (2012) Seven-year trends of CO2 exchange in a tundra ecosystem affected by long-term permafrost thaw. Journal of Geophysical Research-Biogeosciences 117: G2.
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Read PublicationArctic warming has led to permafrost degradation and ground subsidence, created as a result of ground ice melting. Frozen soil organic matter that thaws can increase carbon (C) emissions to the atmosphere, but this can be offset in part by increases in plant growth. The balance of plant and microbial processes, and how this balance changes through time, determines how permafrost ecosystems influence future climate change via the C cycle. This study addressed this question both on short (interannual) and longer (decadal) time periods by measuring C fluxes over a seven-year period at three sites representing a gradient of time since permafrost thaw. All three sites were upland tundra ecosystems located in Interior Alaska but differed in the extent of permafrost thaw and ground subsidence. Results showed an increasing growing season (May – September) trend in gross primary productivity (GPP), net ecosystem exchange (NEE), aboveground net primary productivity (ANPP), and annual NEE at all sites over the seven year study period from 2004 to 2010, but no change in annual and growing season ecosystem respiration (R<sub>eco</sub>). These trends appeared to most closely follow increases in the depth to permafrost that occurred over the same time period. During the seven-year period, sites with more permafrost degradation had significantly greater GPP compared to where degradation was least, but also greater growing season R<sub>eco</sub>. Adding in winter R<sub>eco</sub> decreased, in part, the summer C sink and left only the site with the most permafrost degradation C neutral, with the other sites still C sinks. Annual C balance was strongly dependent on winter R<sub>eco</sub>, which, compared to the growing season, was relatively data-poor due to extreme environmental conditions. As a result, we cannot yet conclude whether the increased NEE in the growing season is truly sustained on an annual basis. If it turns out that winter measurements shown here are an underestimate, we may indeed find these systems are already losing net C to the atmosphere.
Wu Z, Dijkstra P, Koch GW, Hungate BA (2012) Biogeochemical and ecological feedbacks in grassland responses to warming. Nature Climate Change 2(6): 458-461.
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Read PublicationPlant growth often responds rapidly to experimentally simulated climate change<sup><a id="ref-link-1" title="Rustad, L. E. et al. A meta-analysis of the response of soil respiration, net nitrogen mineralization, and aboveground plant growth to experimental ecosystem warming. Oecologia 126, 543-562 (2001)." href="http://www.nature.com/nclimate/journal/v2/n6/full/nclimate1486.html#ref1">1</a>, <a id="ref-link-2" title="Wu, Z., Dijkstra, P., Koch, G. W., Penuelas, J. & Hungate, B. A. Responses of terrestrial ecosystems to temperature and precipitation change: A meta-analysis of experimental manipulation. Glob. Change Biol. 17, 927-942 (2011)." href="http://www.nature.com/nclimate/journal/v2/n6/full/nclimate1486.html#ref2">2</a></sup>. Feedbacks can modulate the initial responses<sup><a id="ref-link-3" title="Harte, J. & Shaw, R. Shifting dominance within a montane vegetation community: Results of a climate-warming experiment. Science 267, 876-880 (1995)." href="http://www.nature.com/nclimate/journal/v2/n6/full/nclimate1486.html#ref3">3</a></sup>, but these feedbacks are difficult to detect when they operate on long timescales<sup><a id="ref-link-4" title="Smith, M. D., Knapp, A. K. & Collins, S. L. A framework for assessing ecosystem dynamics in response to chronic resource alterations induced by global change. Ecology 90, 3279-3289 (2009)." href="http://www.nature.com/nclimate/journal/v2/n6/full/nclimate1486.html#ref4">4</a></sup>. We transplanted intact plant–soil mesocosms down an elevation gradient to expose them to a warmer climate and used collectors and interceptors to simulate changes in precipitation. Here, we show that warming initially increased aboveground net primary productivity in four grassland ecosystems, but the response diminished progressively over nine years. Warming altered the plant community, causing encroachment by species typical of warmer environments and loss of species from the native environment—trends associated with the declining response of plant productivity. Warming stimulated soil nitrogen turnover, which dampened but did not reverse the temporal decline in the productivity response. Warming also enhanced N losses, which may have weakened the expected biogeochemical feedback where warming stimulates N mineralization and plant growth<sup><a id="ref-link-5" title="Rustad, L. E. et al. A meta-analysis of the response of soil respiration, net nitrogen mineralization, and aboveground plant growth to experimental ecosystem warming. Oecologia 126, 543-562 (2001)." href="http://www.nature.com/nclimate/journal/v2/n6/full/nclimate1486.html#ref1">1</a>, <a id="ref-link-6" title="Melillo, J. M. et al. Soil warming, carbon-nitrogen interactions, and forest carbon budgets. Proc. Natl Acad. Sci. USA 108, 9508-9512 (2011)." href="http://www.nature.com/nclimate/journal/v2/n6/full/nclimate1486.html#ref5">5</a>, <a id="ref-link-7" title="Sokolov, A. P. et al. Consequences of considering carbon-nitrogen interactions on the feedbacks between climate and the terrestrial carbon cycle. J. Clim. 21, 3776-3796 (2008)." href="http://www.nature.com/nclimate/journal/v2/n6/full/nclimate1486.html#ref6">6</a></sup>. Our results, describing the responses of four ecosystems to nearly a decade of simulated climate change, indicate that short-term experiments are insufficient to capture the temporal variability and trend of ecosystem responses to environmental change and their modulation through biogeochemical and ecological feedbacks.
2011
Avery LA, Ward DL, and Marks JC (2011) Exercise conditioning decreases downstream movement of pond-reared razorback suckers released into a stream environment. Western North American Naturalist 71(1): 78-85.
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Read PublicationDespite augmentation stocking efforts, wild populations of razorback suckers (Xyrauchen texanus) continue to decline. Endangered razorback suckers are commonly raised in off-channel ponds until maturity (approximately 300 mm TL) and then stocked into the Colorado River or its tributaries. After fish are stocked, they commonly move large distances downstream. We conducted an experiment to determine if downstream dispersal could be reduced through exercise conditioning. Two groups of razorback suckers, exercised and nonexercised, were released into Fossil Creek, Arizona. Prior to release, a subsample from each treatment group was tested in a laboratory flow chamber. Razorback suckers that had been exercise conditioned were able to maintain a position in the flow chamber 2 times longer and at velocities 25 cm · s-1 higher than nonexercised fish. Although the intended method of field data collection via passive-integrated-transponder (PIT) antennas and a remote communication station failed because river otters (Lontra canadensis) preyed upon the released razorback suckers, implanted PIT tags were retained in otter scat. Recovered PIT tags enabled distributional analysis, which indicated that exercised razorback suckers did not move as far downstream from the point of release as nonexercised razorbacks. Exercise conditioning may increase overall fitness of pond-reared razorback suckers, and, consequently, increase the effectiveness of augmentation stocking.
Blankinship JC, Niklaus PA, Hungate BA (2011) A meta-analysis of responses of soil biota to global change. Oecologia 165(3): 553-565.
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Read PublicationGlobal environmental changes are expected to impact the abundance of plants and animals aboveground, but comparably little is known about the responses of belowground organisms. Using meta-analysis, we synthesized results from over 75 manipulative experiments in order to test for patterns in the effects of elevated CO<sub>2</sub>, warming, and altered precipitation on the abundance of soil biota related to taxonomy, body size, feeding habits, ecosystem type, local climate, treatment magnitude and duration, and greenhouse CO<sub>2</sub> enrichment. We found that the positive effect size of elevated CO<sub>2</sub> on the abundance of soil biota diminished with time, whereas the negative effect size of warming and positive effect size of precipitation intensified with time. Trophic group, body size, and experimental approaches best explained the responses of soil biota to elevated CO<sub>2</sub>, whereas local climate and ecosystem type best explained responses to warming and altered precipitation. The abundance of microflora and microfauna, and particularly detritivores, increased with elevated CO<sub>2</sub>, indicative of microbial C limitation under ambient CO<sub>2</sub>. However, the effects of CO<sub>2</sub> were smaller in field studies than in greenhouse studies and were not significant for higher trophic levels. Effects of warming did not depend on taxon or body size, but reduced abundances were more likely to occur at the colder and drier sites. Precipitation limited all taxa and trophic groups, particularly in forest ecosystems. Our meta-analysis suggests that the responses of soil biota to global change are predictable and unique for each global change factor.
Dijkstra P, Blankinship JC, Selmants PC, Hart SC, Koch GW, Schwartz E, Hungate BA (2011) Probing carbon flux patterns through soil microbial metabolic networks using parallel position-specific tracer labeling. Soil Biology and Biochemistry 43(1): 126-132.
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Read PublicationIn order to study controls on metabolic processes in soils, we determined the dynamics of <sup>13</sup>CO<sub>2</sub> production from two position-specific <sup>13</sup>C-labeled pyruvate isotopologues in the presence and absence of glucose, succinate, pine, and legume leaf litter, and under anaerobic conditions. We also compared <sup>13</sup>CO<sub>2</sub> production in soils along a semiarid substrate age gradient in Arizona. We observed that the C from the carboxyl group (C<sub>1</sub>) of pyruvate was lost as CO<sub>2</sub> much faster than its other C atoms (C<sub>2,3</sub>). Addition of glucose, pine and legume leaf litter reduced the ratio between <sup>13</sup>CO<sub>2</sub> production from 1-<sup>13</sup>C pyruvate and 2,3-<sup>13</sup>C pyruvate (C<sub>1</sub>/C<sub>2,3</sub> ratio), whereas anaerobic conditions increased this ratio. Young volcanic soils exhibited a lower C<sub>1</sub>/C<sub>2,3</sub> ratio than older volcanic soils. We interpret a low C<sub>1</sub>/C<sub>2,3</sub> ratio as an indication of increased Krebs cycle activity in response to carbon inputs, while the higher ratio implies a reduced Krebs cycle activity in response to anaerobic conditions. Succinate, a gluconeogenic substrate, reduced <sup>13</sup>CO<sub>2</sub> production from pyruvate to near zero, likely reflecting increased carbohydrate biosynthesis from Krebs cycle intermediates. The difference in <sup>13</sup>CO<sub>2</sub> production rate from pyruvate isotopologues disappeared 4–5 days after pyruvate addition, indicating that C positions were scrambled by ongoing soil microbial transformations. This work demonstrates that metabolic tracers such as pyruvate can be used to determine qualitative aspects of C flux patterns through metabolic pathways of soil microbial communities. Understanding the controls over metabolic processes in soil may improve our understanding of soil C cycling processes.
Dijkstra P, Dalder JJ, Selmants PC, Hart SC, Koch GW, Schwartz E, Hungate BA (2011) Modeling soil metabolic processes using isotopologue pairs of position-specific 13 C-labeled glucose and pyruvate. Soil Biology and Biochemistry 43(9): 1848-1857.
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Read PublicationMost organic carbon (C) in soils eventually turns into CO<sub>2</sub> after passing through microbial metabolic pathways, while providing cells with energy and biosynthetic precursors. Therefore, detailed insight into these metabolic processes may help elucidate mechanisms of soil C cycling processes. Here, we describe a modeling approach to quantify the C flux through metabolic pathways by adding 1-<sup>13</sup>C and 2,3-<sup>13</sup>C pyruvate and 1-<sup>13</sup>C and U-<sup>13</sup>C glucose as metabolic tracers to intact soil microbial communities. The model calculates, assuming steady-state conditions and glucose as the only substrate, the reaction rates through glycolysis, Krebs cycle, pentose phosphate pathway, anaplerotic activity through pyruvate carboxylase, and various biosynthesis reactions. The model assumes a known and constant microbial proportional precursor demand, estimated from literature data. The model is parameterized with experimentally determined ratios of <sup>13</sup>CO<sub>2</sub> production from pyruvate and glucose isotopologue pairs. Model sensitivity analysis shows that metabolic flux patterns are especially responsive to changes in experimentally determined <sup>13</sup>CO<sub>2</sub> ratios from pyruvate and glucose. Calculated fluxes are far less sensitive to assumptions concerning microbial chemical and community composition. The calculated metabolic flux pattern for a young volcanic soil indicates significant pentose phosphate pathway activity in excess of pentose precursor demand and significant anaplerotic activity. These C flux patterns can be used to calculate C use efficiency, energy production and consumption for growth and maintenance purposes, substrate consumption, nitrogen demand, oxygen consumption, and microbial C isotope composition. The metabolic labeling and modeling methods may improve our ability to study the biochemistry and ecophysiology of intact and undisturbed soil microbial communities.
Dijkstra P, Thomas SC, Heinrich PL, Koch GW, Schwartz E, Hungate BA (2011) Effect of temperature on metabolic activity of intact microbial communities: evidence for altered metabolic pathway activity but not for increased maintenance respiration and reduced carbon use efficiency. Soil Biology and Biochemistry 43(10): 2023-2031.
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Read PublicationWe used metabolic tracers and modeling to analyze the response of soil metabolism to a sudden change in temperature from 4 to 20 °C. We hypothesized that intact soil microbial communities would exhibit shifts in pentose phosphate pathway and glycolysis activity in the same way as is regularly observed for individual microorganisms in pure culture. We also hypothesized that increased maintenance respiration at higher temperature would result in greater energy production and reduced carbon use efficiency (CUE). Two hours after temperature increase, respiration increased almost 10-fold. Although all metabolic processes were increased, the relative activity of metabolic processes, biosynthesis, and energy production changed. Pentose phosphate pathway was reduced (17–20%), while activities of specific steps in glycolysis (51%) and Krebs cycle (7–13%) were increased. In contrast, only small but significant changes in biosynthesis (+2%), ATP production (−3%) and CUE (+2%) were observed. In a second experiment, we compared the metabolic responses to temperature increases in soils from high and low elevation. The shift in activity from pentose phosphate pathway to glycolysis with higher temperature was confirmed in both soils, but the responses of Krebs cycle, biosynthesis, ATP production, and CUE were site dependent. Our results indicate that 1) in response to temperature, communities behave biochemically similarly to single species and, 2) our understanding of temperature effects on CUE, energy production and use for maintenance and growth processes is still incomplete.
Dodsworth JA, Hungate BA, Hedlund BP (2011) Ammonia oxidation, denitrification and dissimilatory nitrate reduction to ammonium in two US Great Basin hot springs with abundant ammonia‐oxidizing archaea. Environmental Microbiology 13(8): 2371-2386.
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Read PublicationMany thermophiles catalyse free energy-yielding redox reactions involving nitrogenous compounds; however, little is known about these processes in natural thermal environments. Rates of ammonia oxidation, denitrification and dissimilatory nitrate reduction to ammonium (DNRA) were measured in source water and sediments of two ∼80°C springs in the US Great Basin. Ammonia oxidation and denitrification occurred mainly in sediments. Ammonia oxidation rates measured using <sup>15</sup>N-NO<sub>3</sub><sup>-</sup> pool dilution ranged from 5.5 ± 0.8 to 8.6 ± 0.9 nmol N g<sup>−1</sup> h<sup>−1</sup> and were unaffected or only mildly stimulated by amendment with NH<sub>4</sub>Cl. Denitrification rates measured using acetylene block ranged from 15.8 ± 0.7 to 51 ± 12 nmol N g<sup>−1</sup> h<sup>−1</sup> and were stimulated by amendment with NO<sub>3</sub><sup>-</sup> and complex organic compounds. The DNRA rate in one spring sediment measured using an <sup>15</sup>N-NO<sub>3</sub><sup>-</sup> tracer was 315 ± 48 nmol N g<sup>−1</sup> h<sup>−1</sup>. Both springs harboured distinct planktonic and sediment microbial communities. Close relatives of the autotrophic, ammonia-oxidizing archaeon ‘<em>Candidatus</em> Nitrosocaldus yellowstonii’ represented the most abundant OTU in both spring sediments by 16S rRNA gene pyrotag analysis. Quantitative PCR (qPCR) indicated that ‘<em>Ca</em>. N. yellowstonii’<em>amoA</em> and 16S rRNA genes were present at 3.5–3.9 × 10<sup>8</sup> and 6.4–9.0 × 10<sup>8</sup> copies g<sup>−1</sup> sediment. Potential denitrifiers included members of the <em>Aquificales</em> and <em>Thermales</em>. <em>Thermus</em> spp. comprised < 1% of 16S rRNA gene pyrotags in both sediments and qPCR for <em>T. thermophilus narG</em> revealed sediment populations of 1.3–1.7 × 10<sup>6</sup> copies g<sup>−1</sup> sediment. These data indicate a highly active nitrogen cycle (N-cycle) in these springs and suggest that ammonia oxidation may be a major source of energy fuelling primary production.
Duval BD, Dijkstra P, Natali SM, Megonigal JP, Ketterer ME, Drake BG, Lerdau MT, Gordon G, Anbar AD, Hungate BA (2011) Plant− Soil Distribution of Potentially Toxic Elements in Response to Elevated Atmospheric CO2. Environmental Science & Technology 45(7): 2570-2574.
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Read PublicationThe distribution of contaminant elements within ecosystems is an environmental concern because of these elements’ potential toxicity to animals and plants and their ability to hinder microbial ecosystem services. As with nutrients, contaminants are cycled within and through ecosystems. Elevated atmospheric CO<sub>2</sub> generally increases plant productivity and alters nutrient element cycling, but whether CO<sub>2</sub> causes similar effects on the cycling of contaminant elements is unknown. Here we show that 11 years of experimental CO<sub>2</sub> enrichment in a sandy soil with low organic matter content causes plants to accumulate contaminants in plant biomass, with declines in the extractable contaminant element pools in surface soils. These results indicate that CO<sub>2</sub> alters the distribution of contaminant elements in ecosystems, with plant element accumulation and declining soil availability both likely explained by the CO<sub>2</sub> stimulation of plant biomass. Our results highlight the interdependence of element cycles and the importance of taking a broad view of the periodic table when the effects of global environmental change on ecosystem biogeochemistry are considered.
Fuller BM, Sklar LS, Compson ZG, Adams KJ, Marks JC, Wilcox AC (2011) Ecogeomorphic feedbacks in regrowth of travertine step-pool morphology after dam decommissioning, Fossil Creek, Arizona. Geomorphology 126(3): 314-332.
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The linkages between fluvial geomorphology and aquatic ecosystems are commonly conceptualized as a one-way causal chain in which geomorphic processes create the physical template for ecological dynamics. In streams with a travertine step-pool morphology, however, biotic processes strongly influence the formation and growth of travertine dams, creating the potential for numerous feedbacks. Here we take advantage of the decommissioning of a hydroelectric project on Fossil Creek, Arizona, where restoration of CaCO3-rich baseflow has triggered rapid regrowth of travertine dams, to explore the interactions between biotic and abiotic factors in travertine morphodynamics. We consider three conceptual frameworks, where biotic factors independently modulate the rate of physical and chemical processes that produce travertine dams; combine with abiotic factors in a set of feedback loops; and work in opposition to abiotic processes, such that the travertine step-pool morphology reflects a dynamic balance between dominantly-biotic constructive processes and dominantly-abiotic destructive processes. We consider separately three phases of an idealized life cycle of travertine dams: dam formation, growth, and destruction by erosive floods. Dam formation is catalyzed by abiotic factors (e.g. channel constrictions, and bedrock steps) and biotic factors (e.g. woody debris, and emergent vegetation). From measurements of changes over time in travertine thickness on a bedrock step, we find evidence for a positive feedback between flow hydraulics and travertine accrual. Measurements of organic content in travertine samples from this step show that algal growth contributes substantially to travertine accumulation and suggest that growth is most rapid during seasonal algal blooms. To document vertical growth of travertine dams, we embedded 252 magnets into nascent travertine dams, along a 10 km stretch of river. Growth rates are calculated from changes over time in the magnetic field intensity at the dam surface. At each magnet we record a range of hydraulic and travertine composition variables to characterize the dominant mechanism of growth: abiotic precipitation, algal growth, trapping of organic material, or in situ plant growth. We find: (1) rapid growth of travertine dams following flow restoration, averaging more than 2 cm/year; (2) growth rates decline downstream, consistent with loss of dissolved constituents because of upstream travertine deposition, but also parallel to a decline in organic content in dam surface material and a downstream shift in dominant biotic mechanism; (3) biotic mechanisms are associated with faster growth rates; and (4) correlations between hydraulic attributes and growth rates are more consistent with biotic than abiotic controls. We conclude that the strong influence of living organisms on rates of travertine growth, coupled with the beneficial effects of travertine on ecosystem dynamics, demonstrate a positive feedback between biology and geomorphology. During our two-year study period, erosive flood flows occurred causing widespread removal of travertine. The temporal distribution of travertine growth and erosion over the study period is consistent with a bimodal magnitude– frequency relation in which growth dominates except when large, infrequent storms occur. This model may be useful in other systems where biology exerts strong controls on geomorphic processes.
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Grosse G, Harden J, Turetsky M, McGuire AD, Camill P, Tarnocai C, Frolking S, Schuur EAG, Jorgenson T, Marchenko S, Romanovsky V, Wickland KP, French N, Waldrop M, Bourgeau-Chavez L, Striegl RG (2011) Vulnerability of high-latitude soil organic carbon in North America to disturbance. Journal of Geophysical Research-Biogeosciences 116: G4.
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Read PublicationThis synthesis addresses the vulnerability of the North American high-latitude soil organic carbon (SOC) pool to climate change. Disturbances caused by climate warming in arctic, subarctic, and boreal environments can result in significant redistribution of C among major reservoirs with potential global impacts. We divide the current northern high-latitude SOC pools into (1) near-surface soils where SOC is affected by seasonal freeze-thaw processes and changes in moisture status, and (2) deeper permafrost and peatland strata down to several tens of meters depth where SOC is usually not affected by short-term changes. We address key factors (permafrost, vegetation, hydrology, paleoenvironmental history) and processes (C input, storage, decomposition, and output) responsible for the formation of the large high-latitude SOC pool in North America and highlight how climate-related disturbances could alter this pool's character and size. Press disturbances of relatively slow but persistent nature such as top-down thawing of permafrost, and changes in hydrology, microbiological communities, pedological processes, and vegetation types, as well as pulse disturbances of relatively rapid and local nature such as wildfires and thermokarst, could substantially impact SOC stocks. Ongoing climate warming in the North American high-latitude region could result in crossing environmental thresholds, thereby accelerating press disturbances and increasingly triggering pulse disturbances and eventually affecting the C source/sink net character of northern high-latitude soils. Finally, we assess postdisturbance feedbacks, models, and predictions for the northern high-latitude SOC pool, and discuss data and research gaps to be addressed by future research.
Hedlund BP, McDonald A, Lam J, Dodsworth JA, Brown JR, Hungate BA (2011) Potential role of Thermus thermophilus and T. oshimai in high rates of nitrous oxide (N2O) production in∼ 80° C hot springs in the US Great Basin. Geobiology 9(6): 471-480.
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Read PublicationAmbient nitrous oxide (N<sub>2</sub>O) emissions from Great Boiling Spring (GBS) in the US Great Basin depended on temperature, with the highest flux, 67.8 ± 2.6 μmol N<sub>2</sub>O-N m<sup>−2</sup> day<sup>−1</sup>, occurring in the large source pool at 82 °C. This rate of N<sub>2</sub>O production contrasted with negligible production from nearby soils and was similar to rates from soils and sediments impacted with agricultural fertilizers. To investigate the source of N<sub>2</sub>O, a variety of approaches were used to enrich and isolate heterotrophic micro-organisms, and isolates were screened for nitrate reduction ability. Nitrate-respiring isolates were identified by 16S rRNA gene sequencing as <em>Thermus thermophilus</em> (31 isolates) and <em>T. oshimai</em> (three isolates). All isolates reduced nitrate to N<sub>2</sub>O but not to dinitrogen and were unable to grow with N<sub>2</sub>O as a terminal electron acceptor. Representative <em>T. thermophilus</em> and <em>T. oshimai</em> strains contained genes with 96–98% and 93% DNA identity, respectively, to the nitrate reductase catalytic subunit gene (<em>narG</em>) of <em>T. thermophilus</em> HB8. These data implicate <em>T. thermophilus</em> and <em>T. oshimai</em> in high flux of N<sub>2</sub>O in GBS and raise questions about the genetic basis of the incomplete denitrification pathway in these organisms and on the fate of biogenic N<sub>2</sub>O in geothermal environments.
Johnson KD, Harden J, McGuire AD, Bliss NB, Bockheim JG, Clark M, Nettleton-Hollingsworth T, Jorgenson MT, Kane ES, Mack M, O'Donnell J, Ping C, Schuur EAG, Turetsky MR, Valentine DW (2011) Soil carbon distribution in Alaska in relation to soil-forming factors. Geoderma 167: 71-84.
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Read PublicationThe direction and magnitude of soil organic carbon (SOC) changes in response to climate change remain unclear and depend on the spatial distribution of SOC across landscapes. Uncertainties regarding the fate of SOC are greater in high-latitude systems where data are sparse and the soils are affected by sub-zero temperatures. To address these issues in Alaska, a first-order assessment of data gaps and spatial distributions of SOC was conducted from a recently compiled soil carbon database. Temperature and landform type were the dominant controls on SOC distribution for selected ecoregions. Mean SOC pools (to a depth of 1-m) varied by three, seven and ten-fold across ecoregion, landform, and ecosystem types, respectively. Climate interactions with landform type and SOC were greatest in the uplands. For upland SOC there was a six-fold non-linear increase in SOC with latitude (i.e., temperature) where SOC was lowest in the Intermontane Boreal compared to the Arctic Tundra and Coastal Rainforest. Additionally, in upland systems mineral SOC pools decreased as climate became more continental, suggesting that the lower productivity, higher decomposition rates and fire activity, common in continental climates, interacted to reduce mineral SOC. For lowland systems, in contrast, these interactions and their impacts on SOC were muted or absent making SOC in these environments more comparable across latitudes. Thus, the magnitudes of SOC change across temperature gradients were non-uniform and depended on landform type. Additional factors that appeared to be related to SOC distribution within ecoregions included stand age, aspect, and permafrost presence or absence in black spruce stands. Overall, these results indicate the influence of major interactions between temperature-controlled decomposition and topography on SOC in high-latitude systems. However, there remains a need for more SOC data from wetlands and boreal-region permafrost soils, especially at depths > 1 m in order to fully understand the effects of climate on soil carbon in Alaska.
Lavoie M, Mack MC, Schuur EAG (2011) Effects of elevated nitrogen and temperature on carbon and nitrogen dynamics in Alaskan arctic and boreal soils. Journal of Geophysical Research: Biogeosciences 116: G3.
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Read PublicationPlant productivity in upland tundra and boreal forest is demonstrably limited by nitrogen (N) and indirect evidence from field studies suggests that decomposition by soil microbes may be similarly limited. As climate warms at high latitudes, understanding the response of soil organic matter (SOM) decomposition to increased soil temperature may be crucial for determining the net effect of warming on ecosystem carbon (C) balance because temperature directly affects decomposition but also because it has an indirect effect on C balance via nutrient mineralization. We incubated northern Alaskan soils at two temperatures (5°C and 15°C) and two levels of N addition (with and without) to directly test for N limitation of SOM decomposition and to explore the interaction between temperature and N limitation. Over the entire 924 day incubation of organic and mineral soils from two ecosystem types, we measured microbial respiration; over the initial 90 days of the incubation, we measured microbial biomass N, net N mineralization, and the isotopic signatures (<em>δ</em><sup>13</sup>C and Δ<sup>14</sup>C) of microbial respiration. Across soil layers and ecosystem types, temperature always had a strong positive effect on SOM decomposition rates, whereas N addition had positive, negative, and neutral effects. When C respiration rates were high, the positive N response was generally most strongly expressed, for example, in the organic soils, in the warmer incubation, and at the outset of the experiment. Negative N responses often occurred when C respiration rates were lower, predominantly in mineral soils and at the middle or end of the experiment. In the subset of soil types where we measured the radiocarbon age of respired CO<sub>2</sub>, increased decomposition was related to increased use of older C. Net N mineralization and nitrification were not affected by temperature, but N addition increased net N immobilization in all soil layers and microbial biomass N in organic layers. Our data support the general idea that at least in these high-latitude organic soils, decomposition of labile carbon can be positively stimulated by added N, whereas decomposition of recalcitrant C is suppressed.
Lee H, Schuur EAG, Vogel JG, Lavoie M, Bhadra D, Staudhammer CL (2011) A spatially explicit analysis to extrapolate carbon fluxes in upland tundra where permafrost is thawing. Global Change Biology 17(3): 1379-1393.
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Read PublicationOne of the most important changes in high-latitude ecosystems in response to climatic warming may be the thawing of permafrost soil. In upland tundra, the thawing of ice-rich permafrost can create localized surface subsidence called thermokarst, which may change the soil environment and influence ecosystem carbon release and uptake. We established an intermediate scale (a scale in between point chamber measurements and eddy covariance footprint) ecosystem carbon flux study in Alaskan tundra where permafrost thaw and thermokarst development had been occurring for several decades. The main goal of our study was to examine how dynamic ecosystem carbon fluxes [gross primary production (GPP), ecosystem respiration ( R), and net ecosystem exchange (NEE)] relate to ecosystem variables that incorporate the structural and edaphic changes that co-occur with permafrost thaw and thermokarst development. We then examined how these measured ecosystem carbon fluxes responded to upscaling. For both spatially extensive measurements made intermittently during the peak growing season and intensive measurements made over the entire growing season, ecosystem variables including degree of surface subsidence, thaw depth, and aboveground biomass were selected in a mixed model selection procedure as the 'best' predictors of GPP, R, and NEE. Variables left out of the model (often as a result of autocorrelation) included soil temperature, moisture, and normalized difference vegetation index. These results suggest that the structural changes (surface subsidence, thaw depth, aboveground biomass) that integrate multiple effects of permafrost thaw can be useful components of models used to estimate ecosystem carbon exchange across thermokarst affected landscapes.
Marks JC, Williamson C, Hendrickson DA (2011) Coupling stable isotope studies with food web manipulations to predict the effects of exotic fish: lessons from Cuatro Ciénegas, Mexico. Aquatic Conservation: Marine and Freshwater Ecosystems 21(4): 317-323.
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Read Publication<span class="bullet">1. </span>Exotic species threaten native species worldwide, but their impacts are difficult to predict.
<span class="bullet">2.</span>Stable isotope analysis was combined with field competition experiments to predict how an invasive African cichlid fish, <em>Hemichromis guttatus</em>, might affect native fish in the desert springs of Cuatro Ciénegas, Mexico.
<span class="bullet">3.</span>Stable isotope analysis suggested diet overlap between the invader and juvenile endemic cichlids, and field experiments verified that the invader reduces growth rates of the juvenile endemics through competition, but has smaller effects on adults.
<span class="bullet">4.</span>Competition between juvenile endemic cichlids and the invader was asymmetric, with the exotic out-competing the native, suggesting the potential for competitive exclusion if the invasion is not stopped.
<span class="bullet">5.</span>These results suggest that exotic removal programmes in Cuatro Ciénegas should focus on removing/reducing populations of the exotic cichlid in habitats where juvenile native cichlids are concentrated.
<span class="bullet">6.</span>This approach could help focus efforts to manage exotic species before populations of native species have crashed, when it is too late to intervene.
Natali SM, Schuur EAG, Trucco C, HICKS PRIES CE, Crummer KG, BARON LOPEZ AF (2011) Effects of experimental warming of air, soil and permafrost on carbon balance in Alaskan tundra. Global Change Biology 17(3): 1394-1407.
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Read PublicationThe carbon (C) storage capacity of northern latitude ecosystems may diminish as warming air temperatures increase permafrost thaw and stimulate decomposition of previously frozen soil organic C. However, warming may also enhance plant growth so that photosynthetic carbon dioxide (CO<sub>2</sub>) uptake may, in part, offset respiratory losses. To determine the effects of air and soil warming on CO<sub>2</sub> exchange in tundra, we established an ecosystem warming experiment – the Carbon in Permafrost Experimental Heating Research (CiPEHR) project – in the northern foothills of the Alaska Range in Interior Alaska. We used snow fences coupled with spring snow removal to increase deep soil temperatures and thaw depth (winter warming) and open-top chambers to increase growing season air temperatures (summer warming). Winter warming increased soil temperature (integrated 5–40 cm depth) by 1.5 °C, which resulted in a 10% increase in growing season thaw depth. Surprisingly, the additional 2 kg of thawed soil C m<sup>−2</sup> in the winter warming plots did not result in significant changes in cumulative growing season respiration, which may have been inhibited by soil saturation at the base of the active layer. In contrast to the limited effects on growing-season C dynamics, winter warming caused drastic changes in winter respiration and altered the annual C balance of this ecosystem by doubling the net loss of CO<sub>2</sub> to the atmosphere. While most changes to the abiotic environment at CiPEHR were driven by winter warming, summer warming effects on plant and soil processes resulted in 20% increases in both gross primary productivity and growing season ecosystem respiration and significantly altered the age and sources of CO<sub>2</sub> respired from this ecosystem. These results demonstrate the vulnerability of organic C stored in near surface permafrost to increasing temperatures and the strong potential for warming tundra to serve as a positive feedback to global climate change.
Niboyet A, Brown JR, Dijkstra P, Blankinship JC, Leadley PW, Le Roux X, Barthes L, Barnard RL, Field CB, Hungate BA (2011) Global change could amplify fire effects on soil greenhouse gas emissions. PLoS One 6(6): e20105.
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<h3>Background</h3>
Little is known about the combined impacts of global environmental changes and ecological disturbances on ecosystem functioning, even though such combined impacts might play critical roles in shaping ecosystem processes that can in turn feed back to climate change, such as soil emissions of greenhouse gases.
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<h3>Methodology/Principal Findings</h3>
We took advantage of an accidental, low-severity wildfire that burned part of a long-term global change experiment to investigate the interactive effects of a fire disturbance and increases in CO<sub>2</sub> concentration, precipitation and nitrogen supply on soil nitrous oxide (N<sub>2</sub>O) emissions in a grassland ecosystem. We examined the responses of soil N<sub>2</sub>O emissions, as well as the responses of the two main microbial processes contributing to soil N<sub>2</sub>O production – nitrification and denitrification – and of their main drivers. We show that the fire disturbance greatly increased soil N<sub>2</sub>O emissions over a three-year period, and that elevated CO<sub>2</sub> and enhanced nitrogen supply amplified fire effects on soil N<sub>2</sub>O emissions: emissions increased by a factor of two with fire alone and by a factor of six under the combined influence of fire, elevated CO<sub>2</sub> and nitrogen. We also provide evidence that this response was caused by increased microbial denitrification, resulting from increased soil moisture and soil carbon and nitrogen availability in the burned and fertilized plots.
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<h3>Conclusions/Significance</h3>
Our results indicate that the combined effects of fire and global environmental changes can exceed their effects in isolation, thereby creating unexpected feedbacks to soil greenhouse gas emissions. These findings highlight the need to further explore the impacts of ecological disturbances on ecosystem functioning in the context of global change if we wish to be able to model future soil greenhouse gas emissions with greater confidence.
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Niboyet A, Le Roux X, Dijkstra P, Hungate BA, Barthes L, Blankinship JC, Brown JR, Field CB, Leadley PW (2011) Testing interactive effects of global environmental changes on soil nitrogen cycling. Ecosphere 2(5): 1-24.
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Read PublicationResponses of soil nitrogen (N) cycling to simultaneous and potentially interacting global environmental changes are uncertain. Here, we investigated the combined effects of elevated CO<sub>2</sub>, warming, increased precipitation and enhanced N supply on soil N cycling in an annual grassland ecosystem as part of the Jasper Ridge Global Change Experiment (CA, USA). This field experiment included four treatments—CO<sub>2</sub>, temperature, precipitation, nitrogen—with two levels per treatment (ambient and elevated), and all their factorial combinations replicated six times. We collected soil samples after 7 and 8 years of treatments, and measured gross rates of N mineralization, N immobilization and nitrification, along with potential rates of ammonia oxidation, nitrite oxidation and denitrification. We also determined the main drivers of these microbial activities (soil ammonium and nitrate concentrations, soil moisture, soil temperature, soil pH, and soil CO<sub>2</sub> efflux, as an indicator of soil heterotrophic activity). We found that gross N mineralization responded to the interactive effects of the CO<sub>2</sub>, precipitation and N treatments: N addition increased gross N mineralization when CO<sub>2</sub> and precipitation were either both at ambient or both at elevated levels. However, we found limited evidence for interactions among elevated CO<sub>2</sub>, warming, increased precipitation, and enhanced N supply on the other N cycling processes examined: statistically significant interactions, when found, tended not to persist across multiple dates. Soil N cycling responded mainly to single-factor effects: long-term N addition increased gross N immobilization, potential ammonia oxidation and potential denitrification, while increased precipitation depressed potential nitrite oxidation and increased potential ammonia oxidation and potential denitrification. In contrast, elevated CO<sub>2</sub> and modest warming did not significantly affect any of these microbial N transformations. These findings suggest that global change effects on soil N cycling are primarily additive, and therefore generally predictable from single factor studies.
Posada JM, Schuur EAG (2011) Relationships among precipitation regime, nutrient availability, and carbon turnover in tropical rain forests. Oecologia 165(3): 783-795.
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Read PublicationThe effect of high precipitation regime in tropical forests is poorly known despite indications of its potentially negative effects on nutrient availability and carbon (C) cycling. Our goal was to determine if there was an effect of high rainfall on nitrogen (N) and phosphorous (P) availability and indexes of C cycling in lowland tropical rain forests exposed to a broad range of mean annual precipitation (MAP). We predicted that C turnover time would increase with MAP while the availability of N and P would decrease. We studied seven Neotropical lowland forests covering a MAP range between 2,700 and 9,500 mm. We used radiocarbon (∆<sup>14</sup>C) from the atmosphere and respired from soil organic matter to estimate residence time of C in plants and soils. We also used C, N, and P concentrations and the stable isotope ratio of N (δ<sup>15</sup>N) in live and dead plant tissues and in soils as proxies for nutrient availability. Negative δ<sup>15</sup>N values indicated that the wettest forests had N cycles that did not exhibit isotope-fractionating losses and were potentially N-limited. Element ratios (N:P and C:P) in senescent leaves, litter, and live roots showed that P resorption increased considerably with MAP, which points towards increasing P-limitation under high MAP regimes. Soil C content increased with MAP but C turnover time only showed a weak relationship with MAP, probably due to variations in soil parent material and age along the MAP gradient. In contrast, comparing C turnover directly to nutrient availability showed strong relationships between C turnover time, N availability (δ<sup>15</sup>N), and P availability (N:P) in senescent leaves and litter. Thus, an effect of MAP on carbon cycling appeared to be indirectly mediated by nutrient availability. Our results suggest that soil nutrient availability plays a central role in the dynamic of C cycling in tropical rain forests.
Schirrmeister L, Grosse G, Wetterich S, Overduin PP, Strauss J, Schuur EAG, Hubberten HW (2011) Fossil organic matter characteristics in permafrost deposits of the northeast Siberian Arctic. Journal of Geophysical Research: Biogeosciences 116: G2.
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Read PublicationPermafrost deposits constitute a large organic carbon pool highly vulnerable to degradation and potential carbon release due to global warming. Permafrost sections along coastal and river bank exposures in NE Siberia were studied for organic matter (OM) characteristics and ice content. OM stored in Quaternary permafrost grew, accumulated, froze, partly decomposed, and refroze under different periglacial environments, reflected in specific biogeochemical and cryolithological features. OM in permafrost is represented by twigs, leaves, peat, grass roots, and plant detritus. The vertical distribution of total organic carbon (TOC) in exposures varies from 0.1 wt % of the dry sediment in fluvial deposits to 45 wt % in Holocene peats. Variations in OM parameters are related to changes in vegetation, bioproductivity, pedogenic processes, decomposition, and sedimentation rates during past climate variations. High TOC, high C/N, and low <em>δ</em><sup>13</sup>C reflect less decomposed OM accumulated under wet, anaerobic soil conditions characteristic of interglacial and interstadial periods. Glacial and stadial periods are characterized by less variable, low TOC, low C/N, and high <em>δ</em><sup>13</sup>C values indicating stable environments with reduced bioproductivity and stronger OM decomposition under dryer, aerobic soil conditions. Based on TOC data and updated information on bulk densities, we estimate average organic carbon inventories for ten different stratigraphic units in northeast Siberia, ranging from 7.2 kg C m<sup>−3</sup> for Early Weichselian fluvial deposits, to 33.2 kg C m<sup>−3</sup> for Middle Weichselian Ice Complex deposits, to 74.7 kg C m<sup>−3</sup> for Holocene peaty deposits. The resulting landscape average is likely about 25% lower than previously published permafrost carbon inventories.
Sullivan BW, Kolb TE, Hart SC, Kaye JP, Hungate BA, Dore S, Montes-Helu M (2011) Wildfire reduces carbon dioxide efflux and increases methane uptake in ponderosa pine forest soils of the southwestern USA. Biogeochemistry 104(1-3): 251-265.
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Read PublicationSevere wildfire may cause long-term changes in the soil-atmosphere exchange of carbon dioxide and methane, two gases known to force atmospheric warming. We examined the effect of a severe wildfire 10 years after burning to determine decadal-scale changes in soil gas fluxes following fire, and explored mechanisms responsible for these dynamics. We compared soil carbon dioxide efflux, methane uptake, soil temperature, soil water content, soil O horizon mass, fine root mass, and microbial biomass between a burned site and an unburned site that had similar stand conditions to the burned site before the fire. Compared to the unburned site, soil carbon dioxide efflux was 40% lower and methane uptake was 49% higher at the burned site over the 427-day measurement period. Soil O horizon mass, microbial biomass, fine root mass, and surface soil water content were lower at the burned site than the unburned site, but soil temperature was higher. A regression model showed soil carbon dioxide efflux was more sensitive to changes in soil temperature at the burned site than the unburned site. The relative importance of methane uptake to carbon dioxide efflux was higher at the burned site than the unburned site, but methane uptake compensated for only 1.5% of the warming potential of soil carbon dioxide efflux at the burned site. Our results suggest there was less carbon available at the burned site for respiration by plants and microbes, and the loss of the soil O horizon increased methane uptake in soil at the burned site.
Van Groenigen KJ, Osenberg CW, Hungate BA (2011) Increased soil emissions of potent greenhouse gases under increased atmospheric CO2. Nature 475(7355): 214-216.
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Read PublicationIncreasing concentrations of atmospheric carbon dioxide (CO<sub>2</sub>) can affect biotic and abiotic conditions in soil, such as microbial activity and water content<sup><a id="ref-link-1" title="Zak, D. R., Pregitzer, K. S., King, J. S. & Holmes, W. E. Elevated atmospheric CO2, fine roots and the response of soil microorganisms: a review and hypothesis. New Phytol. 147, 201-222 (2000)" href="http://www.nature.com/nature/journal/v475/n7355/full/nature10176.html#ref1">1</a>, <a id="ref-link-2" title="Pendall, E. et al. Below-ground process responses to elevated CO2 and temperature: a discussion of observations, measurement methods, and models. New Phytol. 162, 311-322 (2004)" href="http://www.nature.com/nature/journal/v475/n7355/full/nature10176.html#ref2">2</a></sup>. In turn, these changes might be expected to alter the production and consumption of the important greenhouse gases nitrous oxide (N<sub>2</sub>O) and methane (CH<sub>4</sub>) (refs <a id="ref-link-3" title="Pendall, E. et al. Below-ground process responses to elevated CO2 and temperature: a discussion of observations, measurement methods, and models. New Phytol. 162, 311-322 (2004)" href="http://www.nature.com/nature/journal/v475/n7355/full/nature10176.html#ref2">2</a>, <a id="ref-link-4" title="Smith, K. A. et al. Exchange of greenhouse gases between soil and atmosphere: interactions of soil physical factors and biological processes. Eur. J. Soil Sci. 54, 779-791 (2003)" href="http://www.nature.com/nature/journal/v475/n7355/full/nature10176.html#ref3">3</a>). However, studies on fluxes of N<sub>2</sub>O and CH<sub>4</sub> from soil under increased atmospheric CO<sub>2</sub> have not been quantitatively synthesized. Here we show, using meta-analysis, that increased CO<sub>2</sub> (ranging from 463 to 780 parts per million by volume) stimulates both N<sub>2</sub>O emissions from upland soils and CH<sub>4</sub> emissions from rice paddies and natural wetlands. Because enhanced greenhouse-gas emissions add to the radiative forcing of terrestrial ecosystems, these emissions are expected to negate at least 16.6 per cent of the climate change mitigation potential previously predicted from an increase in the terrestrial carbon sink under increased atmospheric CO<sub>2</sub> concentrations<sup><a id="ref-link-5" title="Thornton, P. E., Lamarque, J.-F., Rosenbloom, N. A. & Mahowald, N. M. Influence of carbon-nitrogen cycle coupling on land model response to CO2 fertilization and climate variability. Glob. Biogeochem. Cycles 21 GB4018 10.1029/2006GB002868 (2007)" href="http://www.nature.com/nature/journal/v475/n7355/full/nature10176.html#ref4">4</a></sup>. Our results therefore suggest that the capacity of land ecosystems to slow climate warming has been overestimated.
Weber CF, Zak DR, Hungate BA, Jackson RB, Vilgalys R, Evans RD, Schadt CW, Megonigal JP, Kuske CR (2011) Responses of soil cellulolytic fungal communities to elevated atmospheric CO2 are complex and variable across five ecosystems. Environmental microbiology 13(10): 2778-2793.
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Read PublicationElevated atmospheric CO<sub>2</sub> generally increases plant productivity and subsequently increases the availability of cellulose in soil to microbial decomposers. As key cellulose degraders, soil fungi are likely to be one of the most impacted and responsive microbial groups to elevated atmospheric CO<sub>2</sub>. To investigate the impacts of ecosystem type and elevated atmospheric CO<sub>2</sub> on cellulolytic fungal communities, we sequenced 10 677 <em>cbhI</em> gene fragments encoding the catalytic subunit of cellobiohydrolase I, across five distinct terrestrial ecosystem experiments after a decade of exposure to elevated CO<sub>2</sub>. The <em>cbhI</em> composition of each ecosystem was distinct, as supported by weighted Unifrac analyses (all <em>P</em>-values; < 0.001), with few operational taxonomic units (OTUs) being shared across ecosystems. Using a 114-member <em>cbhI</em> sequence database compiled from known fungi, less than 1% of the environmental sequences could be classified at the family level indicating that cellulolytic fungi <em>in situ</em> are likely dominated by novel fungi or known fungi that are not yet recognized as cellulose degraders. Shifts in fungal <em>cbhI</em> composition and richness that were correlated with elevated CO<sub>2</sub> exposure varied across the ecosystems. In aspen plantation and desert creosote bush soils, <em>cbhI</em> gene richness was significantly higher after exposure to elevated CO<sub>2</sub> (550 µmol mol<sup>−1</sup>) than under ambient CO<sub>2</sub> (360 µmol mol<sup>−1</sup> CO<sub>2</sub>). In contrast, while the richness was not altered, the relative abundance of dominant OTUs in desert soil crusts was significantly shifted. This suggests that responses are complex, vary across different ecosystems and, in at least one case, are OTU-specific. Collectively, our results document the complexity of cellulolytic fungal communities in multiple terrestrial ecosystems and the variability of their responses to long-term exposure to elevated atmospheric CO<sub>2</sub>.
Wolken JM, Hollingsworth TN, Rupp TS, Chapin FS, Trainor SF, Barrett TM, Sullivan PF, McGuire AD, Euskirchen ES, Hennon PE, Beever EA, Conn JS, Crone LK, D'Amore DV, Fresco N, Hanley TA, Kielland K, Kruse JJ, Patterson T, Schuur EAG, Verbyla DL, Yarie J (2011) Evidence and implications of recent and projected climate change in Alaska’s forest ecosystems. Ecosphere 2(11): 1-35.
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Read PublicationThe structure and function of Alaska's forests have changed significantly in response to a changing climate, including alterations in species composition and climate feedbacks (e.g., carbon, radiation budgets) that have important regional societal consequences and human feedbacks to forest ecosystems. In this paper we present the first comprehensive synthesis of climate-change impacts on all forested ecosystems of Alaska, highlighting changes in the most critical biophysical factors of each region. We developed a conceptual framework describing climate drivers, biophysical factors and types of change to illustrate how the biophysical and social subsystems of Alaskan forests interact and respond directly and indirectly to a changing climate. We then identify the regional and global implications to the climate system and associated socio-economic impacts, as presented in the current literature. Projections of temperature and precipitation suggest wildfire will continue to be the dominant biophysical factor in the Interior-boreal forest, leading to shifts from conifer- to deciduous-dominated forests. Based on existing research, projected increases in temperature in the Southcentral- and Kenai-boreal forests will likely increase the frequency and severity of insect outbreaks and associated wildfires, and increase the probability of establishment by invasive plant species. In the Coastal-temperate forest region snow and ice is regarded as the dominant biophysical factor. With continued warming, hydrologic changes related to more rapidly melting glaciers and rising elevation of the winter snowline will alter discharge in many rivers, which will have important consequences for terrestrial and marine ecosystem productivity. These climate-related changes will affect plant species distribution and wildlife habitat, which have regional societal consequences, and trace-gas emissions and radiation budgets, which are globally important. Our conceptual framework facilitates assessment of current and future consequences of a changing climate, emphasizes regional differences in biophysical factors, and points to linkages that may exist but that currently lack supporting research. The framework also serves as a visual tool for resource managers and policy makers to develop regional and global management strategies and to inform policies related to climate mitigation and adaptation.
Wu Z, Dijkstra P, Koch GW, Peñuelas J, Hungate BA (2011) Responses of terrestrial ecosystems to temperature and precipitation change: a meta‐analysis of experimental manipulation. Global Change Biology 17(12): 927-942.
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Read PublicationGlobal mean temperature is predicted to increase by 2–7 °C and precipitation to change across the globe by the end of this century. To quantify climate effects on ecosystem processes, a number of climate change experiments have been established around the world in various ecosystems. Despite these efforts, general responses of terrestrial ecosystems to changes in temperature and precipitation, and especially to their combined effects, remain unclear. We used meta-analysis to synthesize ecosystem-level responses to warming, altered precipitation, and their combination. We focused on plant growth and ecosystem carbon (C) balance, including biomass, net primary production (NPP), respiration, net ecosystem exchange (NEE), and ecosystem photosynthesis, synthesizing results from 85 studies. We found that experimental warming and increased precipitation generally stimulated plant growth and ecosystem C fluxes, whereas decreased precipitation had the opposite effects. For example, warming significantly stimulated total NPP, increased ecosystem photosynthesis, and ecosystem respiration. Experimentally reduced precipitation suppressed aboveground NPP (ANPP) and NEE, whereas supplemental precipitation enhanced ANPP and NEE. Plant productivity and ecosystem C fluxes generally showed higher sensitivities to increased precipitation than to decreased precipitation. Interactive effects of warming and altered precipitation tended to be smaller than expected from additive, single-factor effects, though low statistical power limits the strength of these conclusions. New experiments with combined temperature and precipitation manipulations are needed to conclusively determine the importance of temperature–precipitation interactions on the C balance of terrestrial ecosystems under future climate conditions.
Wu Z, Koch GW, Dijkstra P, Bowker MA, Hungate BA (2011) Responses of ecosystem carbon cycling to climate change treatments along an elevation gradient. Ecosystems 14(7): 1066-1080.
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Read PublicationGlobal temperature increases and precipitation changes are both expected to alter ecosystem carbon (C) cycling. We tested responses of ecosystem C cycling to simulated climate change using field manipulations of temperature and precipitation across a range of grass-dominated ecosystems along an elevation gradient in northern Arizona. In 2002, we transplanted intact plant–soil mesocosms to simulate warming and used passive interceptors and collectors to manipulate precipitation. We measured daytime ecosystem respiration (ER) and net ecosystem C exchange throughout the growing season in 2008 and 2009. Warming generally stimulated ER and photosynthesis, but had variable effects on daytime net C exchange. Increased precipitation stimulated ecosystem C cycling only in the driest ecosystem at the lowest elevation, whereas decreased precipitation showed no effects on ecosystem C cycling across all ecosystems. No significant interaction between temperature and precipitation treatments was observed. Structural equation modeling revealed that in the wetter-than-average year of 2008, changes in ecosystem C cycling were more strongly affected by warming-induced reduction in soil moisture than by altered precipitation. In contrast, during the drier year of 2009, warming induced increase in soil temperature rather than changes in soil moisture determined ecosystem C cycling. Our findings suggest that warming exerted the strongest influence on ecosystem C cycling in both years, by modulating soil moisture in the wet year and soil temperature in the dry year.
2010
Anderson RG, Canadell JG, Randerson JT, Jackson RB, Hungate BA, Baldocchi DD, Ban-Weiss GA, Bonan GB, Caldeira K, Cao L, Diffenbaugh NS, Gurney KR, Kueppers LM, Law BE, Luyssaert S, O'Halloran TL (2010) Biophysical considerations in forestry for climate protection. Frontiers in Ecology and the Environment 9(3): 174-182.
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Read PublicationForestry – including afforestation (the planting of trees on land where they have not recently existed), reforestation, avoided deforestation, and forest management – can lead to increased sequestration of atmospheric carbon dioxide and has therefore been proposed as a strategy to mitigate climate change. However, forestry also influences land-surface properties, including albedo (the fraction of incident sunlight reflected back to space), surface roughness, and evapotranspiration, all of which affect the amount and forms of energy transfer to the atmosphere. In some circumstances, these biophysical feedbacks can result in local climate warming, thereby counteracting the effects of carbon sequestration on global mean temperature and reducing or eliminating the net value of climate-change mitigation projects. Here, we review published and emerging research that suggests ways in which forestry projects can counteract the consequences associated with biophysical interactions, and highlight knowledge gaps in managing forests for climate protection. We also outline several ways in which biophysical effects can be incorporated into frameworks that use the maintenance of forests as a climate protection strategy.
Billings SA, Lichter J, Ziegler SE, Hungate BA, de B Richter D (2010) A call to investigate drivers of soil organic matter retention vs. mineralization in a high CO 2 world. Soil Biology and Biochemistry 42(4): 665-668.
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Read PublicationUnderstanding how elevated atmospheric CO<sub>2</sub> alters the formation and decomposition of soil organic carbon (SOC) is important but challenging. If elevated CO<sub>2</sub> induces even small changes in rates of formation or decay of SOC, there could be substantial feedbacks on the atmosphere's concentration of CO<sub>2</sub>. However, the long turnover times of many SOC pools – decades to centuries – make the detection of changes in the soil's pool size difficult. Long-term CO<sub>2</sub> enrichment experiments have offered unprecedented opportunities to explore these issues in intact ecosystems for more than a decade. Increased NPP with elevated CO<sub>2</sub> has prompted the hypothesis that SOC may increase at the same time that increased vegetation nitrogen (N) uptake and accumulation indicates probable declines in SON. Varying investigators thus have hypothesized that SOC will increase and SON will decline to explain increased NPP with elevated CO<sub>2</sub>; researchers also invoke biogeochemical theory and stoichiometric constraints to argue for strong limitations on the co-occurrence of these phenomena. We call for researchers to investigate two broad research questions to elucidate the drivers of these processes. First, we ask how elevated CO<sub>2</sub> influences compound structure and stoichiometry of that proportion of NPP retained by soil profiles for relatively long time periods. We also call for investigations of the mechanisms underlying the decomposition of mineralizable organic matter with elevated CO<sub>2</sub>. Specifically, we need to understand how elevated CO<sub>2</sub> influences microbial priming (driven by enhanced microbial energy needs associated with increases in biomass or activity) and microbial mining of N (driven by enhanced microbial N demand associated with greater vegetative N uptake), two processes that necessarily will be constrained by the stoichiometry of both substrates and microbial demands. Applying technologies such as nuclear magnetic resonance and the detection of biomarkers that reveal organic matter structure and origins, and studying microbial stoichiometric constraints, will dramatically improve our ability to predict future patterns of ecosystem C and N cycling.
Blankinship JC, Brown JR, Dijkstra P, Allwright MC, Hungate BA (2010) Response of terrestrial CH4 uptake to interactive changes in precipitation and temperature along a climatic gradient. Ecosystems 13(8): 1157-1170.
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Read PublicationWe determined the response of terrestrial methane (CH<sub>4</sub>) uptake to 4 years of full-factorial manipulations of precipitation and temperature in four ecosystems along a 50 km warm and dry to cold and wet climatic gradient (desert grassland, pinyon-juniper woodland, ponderosa pine forest, and mixed conifer forest). Our goals were to determine whether ecosystem-specific, intraannual, and interactive responses to altered precipitation and warming are quantitatively important. Passive collectors and interceptors increased (+50% per event) and reduced (−30% per event) the quantity of precipitation delivered to experimental plant–soil mesocosms, and downward transfer along the elevation gradient warmed mesocosms by 1.8°C on average. Methane uptake in the colder and wetter ecosystems along the gradient decreased with increasing precipitation, especially during the wet season. The warmer and drier ecosystems, however, responded more strongly to warming, exhibiting less CH<sub>4</sub> uptake with increasing temperature. We found no interaction between altered precipitation and warming in any ecosystem. Soil CH<sub>4</sub> consumption in the laboratory was a strong predictor of ecosystem differences in field CH<sub>4</sub> consumption, but was a poor predictor of the effects of climatic change observed in the field. Based on our results, future climate scenarios that are wet and warm will cause the largest reduction in terrestrial CH<sub>4</sub> uptake across ecosystem types.
Blankinship JC, Brown JR, Dijkstra P, Hungate BA (2010) Effects of interactive global changes on methane uptake in an annual grassland. Journal of Geophysical Research: Biogeosciences 115: G2.
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Read Publication<span class="paraNumber">[1]</span> The future size of the terrestrial methane (CH<sub>4</sub>) sink of upland soils remains uncertain, along with potential feedbacks to global warming. Much of the uncertainty lies in our lack of knowledge about potential interactive effects of multiple simultaneous global environmental changes. Field CH<sub>4</sub> fluxes and laboratory soil CH<sub>4</sub> consumption were measured five times during 3 consecutive years in a California annual grassland exposed to 8 years of the full factorial combination of ambient and elevated levels of precipitation, temperature, atmospheric CO<sub>2</sub> concentration, and N deposition. Across all sampling dates and treatments, increased precipitation caused a 61% reduction in field CH<sub>4</sub> uptake. However, this reduction depended quantitatively on other global change factors. Higher precipitation reduced CH<sub>4</sub> uptake when temperature or N deposition (but not both) increased, and under elevated CO<sub>2</sub> but only late in the growing season. Warming alone also decreased CH<sub>4</sub> uptake early in the growing season, which was partly explained by a decrease in laboratory soil CH<sub>4</sub> consumption. Atmospheric CH<sub>4</sub> models likely need to incorporate nonadditive interactions, seasonal interactions, and interactions between methanotrophy and methanogenesis. Despite the complexity of interactions we observed in this multifactor experiment, the outcome agrees with results from single-factor experiments: an increased terrestrial CH<sub>4</sub> sink appears less likely than a reduced one.
Boby LA, Schuur EAG, Mack MC, Verbyla D, Johnstone JF (2010) Quantifying fire severity, carbon, and nitrogen emissions in Alaska’s boreal forest. Ecological Applications 20(6): 1633-1647.
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Read PublicationThe boreal region stores a large proportion of the world's terrestrial carbon (C) and is subject to high-intensity, stand-replacing wildfires that release C and nitrogen (N) stored in biomass and soils through combustion. While severity and extent of fires drives overall emissions, methods for accurately estimating fire severity are poorly tested in this unique region where organic soil combustion is responsible for a large proportion of total emissions. We tested a method using adventitious roots on black spruce trees (<span class="genusSpeciesInfoAsset">Picea mariana</span>) in combination with canopy allometry to reconstruct prefire organic soil layers and canopy biomass in boreal black spruce forests of Alaska (USA), thus providing a basis for more accurately quantifying fire severity levels. We calibrated this adventitious-root-height method in unburned spruce stands and then tested it by comparing our biomass and soils estimates reconstructed in burned stands with actual prefire stand measurements. We applied this approach to 38 black spruce stands burned in 2004 in Alaska, where we measured organic soil and stand characteristics and estimated the amount of soil and canopy biomass, as well as C and N pools, consumed by fire. These high-intensity quantitative estimates of severity were significantly correlated to a semiquantitative visual rapid assessment tool, the composite burn index (CBI). This index has proved useful for assessing fire severity in forests in the western United States but has not yet been widely tested in the boreal forest. From our study, we conclude that using postfire measurements of adventitious roots on black spruce trees in combination with soils and tree data can be used to reconstruct prefire organic soil depths and biomass pools, providing accurate estimates of fire severity and emissions. Furthermore, using our quantitative reconstruction we show that CBI is a reasonably good predictor of biomass and soil C loss at these sites, and it shows promise for rapidly estimating fire severity across a wide range of boreal black spruce forest types, especially where the use of high-intensity measurements may be limited by cost and time.
Campbell BJ, Polson SW, Hanson TE, Mack MC, Schuur EAG (2010) The effect of nutrient deposition on bacterial communities in Arctic tundra soil. Environmental Microbiology 12(7): 1842-1854.
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Read PublicationThe microbial communities of high-latitude ecosystems are expected to experience rapid changes over the next century due to climate warming and increased deposition of reactive nitrogen, changes that will likely affect microbial community structure and function. In moist acidic tundra (MAT) soils on the North Slope of the Brooks Range, Alaska, substantial losses of C and N were previously observed after long-term nutrient additions. To analyse the role of microbial communities in these losses, we utilized 16S rRNA gene tag pyrosequencing coupled with community-level physiological profiling to describe changes in MAT bacterial communities after short- and long-term nutrient fertilization in four sets of paired control and fertilized MAT soil samples. Bacterial diversity was lower in long-term fertilized plots. The <em>Acidobacteria</em> were one of the most abundant phyla in all soils and distinct differences were noted in the distributions of<em>Acidobacteria</em> subgroups between mineral and organic soil layers that were also affected by fertilization. In addition, <em>Alpha</em>- and <em>Gammaproteobacteria</em> were more abundant in long-term fertilized samples compared with control soils. The dramatic increase in sequences within the <em>Gammaproteobacteria</em> identified as <em>Dyella</em> spp. (order<em>Xanthomonadales</em>) in the long-term fertilized samples was confirmed by quantitative PCR (qPCR) in several samples. Long-term fertilization was also correlated with shifts in the utilization of specific substrates by microbes present in the soils. The combined data indicate that long-term fertilization resulted in a significant change in microbial community structure and function linked to changes in carbon and nitrogen availability and shifts in above-ground plant communities.
Dore S, Kolb TE, Montes-Helu M, Eckert SE, Sullivan BW, Hungate BA, Kaye JP, Hart SC, Koch GW, Finkral A (2010) Carbon and water fluxes from ponderosa pine forests disturbed by wildfire and thinning. Ecological Applications 20(3):663-683.
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Read PublicationDisturbances alter ecosystem carbon dynamics, often by reducing carbon uptake and stocks. We compared the impact of two types of disturbances that represent the most likely future conditions of currently dense ponderosa pine forests of the southwestern United States: (1) high-intensity fire and (2) thinning, designed to reduce fire intensity. High-severity fire had a larger impact on ecosystem carbon uptake and storage than thinning. Total ecosystem carbon was 42% lower at the intensely burned site, 10 years after burning, than at the undisturbed site. Eddy covariance measurements over two years showed that the burned site was a net annual source of carbon to the atmosphere whereas the undisturbed site was a sink. Net primary production (NPP), evapotranspiration (ET), and water use efficiency were lower at the burned site than at the undisturbed site. In contrast, thinning decreased total ecosystem carbon by 18%, and changed the site from a carbon sink to a source in the first post-treatment year. Thinning also decreased ET, reduced the limitation of drought on carbon uptake during summer, and did not change water use efficiency. Both disturbances reduced ecosystem carbon uptake by decreasing gross primary production (55% by burning, 30% by thinning) more than total ecosystem respiration (TER; 33–47% by burning, 18% by thinning), and increased the contribution of soil carbon dioxide efflux to TER. The relationship between TER and temperature was not affected by either disturbance. Efforts to accurately estimate regional carbon budgets should consider impacts on carbon dynamics of both large disturbances, such as high-intensity fire, and the partial disturbance of thinning that is often used to prevent intense burning. Our results show that thinned forests of ponderosa pine in the southwestern United States are a desirable alternative to intensively burned forests to maintain carbon stocks and primary production.
Fuller BM, Sklar LS, Compson Z, Adams KJ, Marks JC, and Wilcox AC (2010) Ecogeomorphic feedbacks in regrowth of travertine step-pool morphology after dam decomissioning, Fossil Creek, Arizona. Geomorphology 126(3-4): 314-332.
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Read PublicationThe linkages between fluvial geomorphology and aquatic ecosystems are commonly conceptualized as a one-way causal chain inwhich geomorphic processes create the physical template for ecological dynamics. In streams with a travertine step-pool morphology, however, biotic processes strongly influence the formation and growth of travertinedams, creating the potential fornumerous feedbacks.Herewetakeadvantage of the decommissioning of a hydroelectricproject on Fossil Creek,Arizona,where restoration of CaCO3-rich baseflowhas triggered rapid regrowth of travertine dams, to explore the interactions between biotic and abiotic factors in travertine morphodynamics. We consider three conceptual frameworks, where biotic factors independently modulate the rate of physical and chemical processes that produce travertine dams; combinewith abiotic factors in a set of feedback loops; handwork in opposition to abiotic processes, such that the travertine step-pool morphology reflects a dynamic balance between dominantly-biotic constructive processes and dominantly-abiotic destructive processes. We consider separately three phases of an idealized life cycle of travertine dams: dam formation, growth, and destruction by erosive floods. Dam formation is catalyzed by abiotic factors (e.g. channel constrictions, and bedrock steps) and biotic factors (e.g. woody debris, and emergent vegetation). From measurements of changes over time in travertine thickness on a bedrock step, we find evidence for a positive feedback between flow hydraulics and travertine accrual. Measurements of organic content in travertine samples from this step show that algal growth contributes substantially to travertine accumulation and suggest that growth is most rapid during seasonal algal blooms. To document vertical growth of travertine dams, we embedded 252 magnets into nascent travertine dams, along a 10 km stretch of river. Growth rates are calculated from changes over time in themagnetic field intensity at the dam surface. At each magnet we record a range of hydraulic and travertine composition variables to characterize the dominant mechanism of growth: abiotic precipitation, algal growth, trapping of organic material, or in situ plant growth.We find: (1) rapid growth of travertine damsfollowingflowrestoration, averagingmore than 2 cm/year; (2) growth rates decline downstream, consistent with loss of dissolved constituents because of upstream travertine deposition, but also parallel to a decline in organic content in dam surface material and a downstream shift in dominant biotic mechanism; (3) biotic mechanisms are associated with faster growth rates; and (4) correlations between hydraulic attributes and growth rates are more consistent with biotic than abiotic controls.We conclude that the strong influence of living organisms on rates of travertine growth, coupled with the beneficial effects of travertine on ecosystem dynamics, demonstrate a positive feedback between biology and geomorphology. During our two-year study period, erosive flood flows occurred causing widespread removal of travertine. The temporal distribution of travertine growth and erosion over the study period is consistent with a bimodal magnitude– frequency relation in which growth dominates except when large, infrequent storms occur. This model may be useful in other systems where biology exerts strong controls on geomorphic processes.
Jorgenson MT, Romanovsky V, Harden J, Shur Y, O'Donnell J, Schuur EAG, Kanevskiy M, Marchenko S (2010) Resilience and vulnerability of permafrost to climate change. Canadian Journal of Forest Research 40(7): 1219-1236.
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Read PublicationThe resilience and vulnerability of permafrost to climate change depends on complex interactions among topography, water, soil, vegetation, and snow, which allow permafrost to persist at mean annual air temperatures (MAATs) as high as +2 °C and degrade at MAATs as low as –20 °C. To assess these interactions, we compiled existing data and tested effects of varying conditions on mean annual surface temperatures (MASTs) and 2 m deep temperatures (MADTs) through modeling. Surface water had the largest effect, with water sediment temperatures being ~10 °C above MAAT. A 50% reduction in snow depth reduces MADT by 2 °C. Elevation changes between 200 and 800 m increases MAAT by up to 2.3 °C and snow depths by ~40%. Aspect caused only a ~1 °C difference in MAST. Covarying vegetation structure, organic matter thickness, soil moisture, and snow depth of terrestrial ecosystems, ranging from barren silt to white spruce (Picea glauca(Moench) Voss) forest to tussock shrub, affect MASTs by ~6 °C and MADTs by ~7 °C. Groundwater at 2–7 °C greatly affects lateral and internal permafrost thawing. Analyses show that vegetation succession provides strong negative feedbacks that make permafrost resilient to even large increases in air temperatures. Surface water, which is affected by topography and ground ice, provides even stronger negative feedbacks that make permafrost vulnerable to thawing even under cold temperatures.
Kuhry P, Dorrepaal E, Hugelius G, Schuur EAG, Tarnocai C (2010) Potential remobilization of belowground permafrost carbon under future global warming. Permafrost and Periglacial Processes 21(2): 208-214.
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Read PublicationResearch on permafrost carbon has dramatically increased in the past few years, A new estimate of 1672 Pg C of belowground organic carbon in the northern circumpolar permafrost region more than doubles the previous value and highlights the potential role of permafrost carbon in the Earth S ystem. Uncertainties in this new estimate remain due to relatively few available pedon data for certain geographic sectors and the deeper cryoturbated soil horizons, and the large polygon size in the soil maps used for upscaling. The large permafrost carbon pool is not equally distributed across the landscape: peat deposits, cryoturbated soils and the loess-like deposits of the yedoma complex contain disproportionately large amounts of soil organic matteL often exhibiting a low degree of decomposition. Recent findings in Alaska and northern S weden provide strong evidence that the deeper soil carbon in permafrost terrain is starting to be released, supporting previous reports from S iberia. The permafrost carbon pool is not yet fully integrated in climate and ecosystem models and an important objective should be to define typical pedons appropriate for model setups. The thawing permafrost carbon feedback needs to be included in model projections of future climate change.
Lee H, Schuur EAG, Vogel JG (2010) Soil CO2 production in upland tundra where permafrost is thawing. Journal of Geophysical Research: Biogeosciences 115: G1.
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Read PublicationPermafrost soils store nearly half of global soil carbon (C), and therefore permafrost thawing could lead to large amounts of greenhouse gas emissions via decomposition of soil organic matter. When ice-rich permafrost thaws, it creates a localized surface subsidence called thermokarst terrain, which changes the soil microenvironment. We used soil profile CO<sub>2</sub> measurements to understand the response of belowground C emissions for different soil depths from upland tundra as a result of permafrost thaw and thermokarst development. We established sites in central Alaska, where permafrost thaw and thermokarst development had been monitored for the past 2 decades. Cumulative growing season CO<sub>2</sub> production averaged for 3 years (2005–2007) ranged from 177 to 270 g CO<sub>2</sub>-C m<sup>−2</sup> and was lowest in the least disturbed moist acidic tundra and highest where thawing of permafrost and thermokarst was most pronounced. We were able to explain 55% of variability in growing season soil CO<sub>2</sub>production using surface subsidence, soil temperature, and site differences. This was likely a direct effect of permafrost thaw and thermokarst development and an indirect effect of changes in microsite soil temperature and surface moisture content, which stimulated soil organic matter decomposition and root respiration. We also observed unusually high CO<sub>2</sub> concentrations in the early growing season, which may be attributable to trapped CO<sub>2</sub> within air pockets in the frozen soil. Taken together, these results supported the projection that permafrost thaw and thermokarst development will increase belowground carbon emissions in the upland tundra.
Marks JC, Haden GA, O’Neill M, Pace C (2010) Effects of flow restoration and exotic species removal on recovery of native fish: lessons from a dam decommissioning. Restoration Ecology 18(6): 934-943.
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Flow diversion and invasive species are two major threats to freshwater ecosystems, threats that restoration efforts attempt to redress. Yet, few restoration projects monitor whether removal of these threats improve target charac- teristics of the ecosystem. Fewer still have an appropriate experimental design from which causal inferences can be drawn as to the relative merits of removing exotic fish, restoring flow, or both. We used a dam decommissioning in Fossil Creek, Arizona, to compare responses of native fish to exotic fish removal and flow restoration, using a before- after-control-impact design with three impact treatments: flow restoration alone where exotics had not been present, flow restoration and exotic fish removal, and flow restora- tion where exotics remain and a control reach that was unaffected by restoration actions. We show that removal of exotic fish dramatically increased native fish abundance.
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Flow restoration also increased native fish abundance, but the effect was smaller than that from removing exotics. Flow restoration had no effect where exotic fish remained, although it may have had other benefits to the ecosystem. The cost to restore flow ($12 million) was considerably higher than that to eradicate exotics ($1.1 million). The long-term influence of flow restoration could increase, as travertine dams grow and re-shape the creek increasing habitat for native fish. But in the 2-year period consid- ered here, the return on investment for extirpating exotics far exceeded that from flow restoration. Projects aimed to restore native fish by restoring flow should also consider the additional investment required to eradicate exotic fish.
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McGuire AD, Macdonald RW, Schuur EAG, Harden JW, Kuhry P, Hayes DJ, Christensen TR, Heimann M (2010) The carbon budget of the northern cryosphere region. Current Opinion in Environmental Sustainability 2(4): 231-236.
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Read PublicationThe northern cryosphere is undergoing substantial warming of permafrost and loss of sea ice. Release of stored carbon to the atmosphere in response to this change has the potential to affect the global climate system. Studies indicate that the northern cryosphere has been not only a substantial sink for atmospheric CO<sub>2</sub> in recent decades, but also an important source of CH<sub>4</sub> because of emissions from wetlands and lakes. Analyses suggest that the sensitivity of the carbon cycle of the region over the 21st Century is potentially large, but highly uncertain because numerous pathways of response will be affected by warming. Further research should focus on sensitive elements of the carbon cycle such as the consequences of increased fire disturbance, permafrost degradation, and sea ice loss in the northern cryosphere region.
Niboyet A, Barthes L, Hungate BA, Le Roux X, Bloor JMG, Ambroise A, Fontaine SF, Price PM, Leadley PW (2010) Responses of soil nitrogen cycling to the interactive effects of elevated CO2 and inorganic N supply. Plant and Soil 327(1): 35-47.
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Read PublicationDespite increasing interest in the effects of climate change on soil processes, the response of nitrification to elevated CO<sub>2</sub>remains unclear. Responses may depend on soil nitrogen (N) status, and inferences may vary depending on the methodological approach used. We investigated the interactive effects of elevated CO<sub>2</sub> and inorganic N supply on gross nitrification (using <sup>15</sup>N pool dilution) and potential nitrification (using nitrifying enzyme activity assays) in <em class="EmphasisTypeItalic ">Dactylis glomerata</em> mesocosms. We measured the responses of putative drivers of nitrification (NH<span class="Stack"><sub>4</sub><sup>+</sup></span> production, NH<span class="Stack"><sub>4</sub><sup>+</sup></span> consumption, and soil environmental conditions) and of potential denitrification, a process functionally linked to nitrification. Gross nitrification was insensitive to all treatments, whereas potential nitrification was higher in the high N treatment and was further stimulated by elevated CO<sub>2</sub> in the high N treatment. Gross mineralization and NH<span class="Stack"><sub>4</sub><sup>+</sup></span> consumption rates were also significantly increased in response to elevated CO<sub>2</sub> in the high N treatment, while potential denitrification showed a significant increase in response to N addition. The discrepancy between the responses of gross and potential nitrification to elevated CO<sub>2</sub> and inorganic N supply suggest that these measurements provide different information, and should be used as complementary approaches to understand nitrification response to global change.
2009
Compson Z, Mier MZ, and Marks JC (2009) Effects of travertine and flow on leaf retention in Fossil Creek, Ariona. Hydrobiologia 630(1): 187-197.
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Read PublicationLeaf retention is important in transferring energy from riparian trees to stream food webs. Retention increases with geomorphic complexity such as substrate coarseness, sinuosity, and the presence of debris dams. High discharge can reduce retention, particularly when streams lack physical trapping features. Travertine formations, caused by calcium carbonate deposition, can alter stream morphology. To date, however, we know of no study testing the effect of travertine on leaf retention. This study capitalized on a river restoration project in Fossil Creek, Arizona, where water was returned to the channel after a century of diversion. We examined how the fixed factors Flow (before and after restoration) and Morphology (travertine and riffle-pool sites) affected leaf retention. Leaf retention was higher in sites where travertine forms barriers across the river, relative to sites with riffle-pool morphology. Most leaves retained in travertine reaches were concentrated at the bottom of pools formed between dams. Although flow restoration did not alter retention rates across all sites, it diminished them at travertine sites, indicating an interaction between stream flow and morphology. We conclude that stream complexity and leaf retention are enhanced by travertine deposition but that high discharge can reduce the retentive capacity of in-stream structures.
Coyle JS, Dijkstra P, Doucett RR, Schwartz E, Hart SC, Hungate BA (2009) Relationships between C and N availability, substrate age, and natural abundance 13 C and 15 N signatures of soil microbial biomass in a semiarid climate. Soil Biology and Biochemistry 41(8): 1605-1611.
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Read PublicationSoil microbial organisms are central to carbon (C) and nitrogen (N) transformations in soils, yet not much is known about the stable isotope composition of these essential regulators of element cycles. We investigated the relationship between C and N availability and stable C and N isotope composition of soil microbial biomass across a three million year old semiarid substrate age gradient in northern Arizona. The <em>δ</em><sup>15</sup>N of soil microbial biomass was on average 7.2‰ higher than that of soil total N for all substrate ages and 1.6‰ higher than that of extractable N, but not significantly different for the youngest and oldest sites. Microbial <sup>15</sup>N enrichment relative to soil extractable and total N was low at the youngest site, increased to a maximum after 55,000 years, and then decreased slightly with age. The degree of <sup>15</sup>N enrichment of microbial biomass correlated negatively with the C:N mass ratio of the soil extractable pool. The <em>δ</em><sup>13</sup>C signature of soil microbial biomass was 1.4‰ and 4.6‰ enriched relative to that of soil total and extractable pools respectively and showed significant differences between sites. However, microbial <sup>13</sup>C enrichment was unrelated to measures of C and N availability. Our results confirm that <sup>15</sup>N, but not <sup>13</sup>C enrichment of soil microbial biomass reflects changes in C and N availability and N processing during long-term ecosystem development.
Craine JM, Elmore AJ, Aidar MPM, Bustamante M, Dawson TE, Hobbie EA, Kahmen A, Mack MC, McLauchlan KK, Michelsen A, Nardoto GB, Pardo LH, Penuelas J, Reich PB, Schuur EAG, Stock WD, Templer PH, Virginia RA, Welker JM, Wright IJ (2009) Global patterns of foliar nitrogen isotopes and their relationships with climate, mycorrhizal fungi, foliar nutrient concentrations, and nitrogen availability. New Phytologist 183(4): 980-992.
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Read PublicationRatios of nitrogen (N) isotopes in leaves could elucidate underlying patterns of N cycling across ecological gradients. To better understand global-scale patterns of N cycling, we compiled data on foliar N isotope ratios (δ<sup>15</sup>N), foliar N concentrations, mycorrhizal type and climate for over 11 000 plants worldwide. Arbuscular mycorrhizal, ectomycorrhizal, and ericoid mycorrhizal plants were depleted in foliar δ<sup>15</sup>N by 2‰, 3.2‰, 5.9‰, respectively, relative to nonmycorrhizal plants. Foliar δ<sup>15</sup>N increased with decreasing mean annual precipitation and with increasing mean annual temperature (MAT) across sites with MAT ≥ −0.5°C, but was invariant with MAT across sites with MAT < −0.5°C. In independent landscape-level to regional-level studies, foliar δ<sup>15</sup>N increased with increasing N availability; at the global scale, foliar δ<sup>15</sup>N increased with increasing foliar N concentrations and decreasing foliar phosphorus (P) concentrations. Together, these results suggest that warm, dry ecosystems have the highest N availability, while plants with high N concentrations, on average, occupy sites with higher N availability than plants with low N concentrations. Global-scale comparisons of other components of the N cycle are still required for better mechanistic understanding of the determinants of variation in foliar δ<sup>15</sup>N and ultimately global patterns in N cycling.
Durham WB, Pathare AV, Stern LA, Lenferink HJ (2009) Mobility of icy sand packs, with application to Martian permafrost. Geophysical Research Letters 36(23).
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Read PublicationThe physical state of water on Mars has fundamental ramifications for both climatology and astrobiology. The widespread presence of “softened” Martian landforms (such as impact craters) can be attributed to viscous creep of subsurface ground ice. We present laboratory experiments designed to determine the minimum amount of ice necessary to mobilize topography within Martian permafrost. Our results show that the jammed-to-mobile transition of icy sand packs neither occurs at fixed ice content nor is dependent on temperature or stress, but instead correlates strongly with the maximum dry packing density of the sand component. Viscosity also changes rapidly near the mobility transition. The results suggest a potentially lower minimum volatile inventory for the impact-pulverized megaregolith of Mars. Furthermore, the long-term preservation of partially relaxed craters implies that the ice content of Martian permafrost has remained close to that at the mobility transition throughout Martian history.
Hungate BA, Van Groenigen KJ, Six J, Jastrow JD, Luo Y, De Graaff M, van Kessel C, Osenberg CW (2009) Assessing the effect of elevated carbon dioxide on soil carbon: a comparison of four meta‐analyses. Global Change Biology 15(8): 2020-2034.
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Read PublicationSoil is the largest reservoir of organic carbon (C) in the terrestrial biosphere and soil C has a relatively long mean residence time. Rising atmospheric carbon dioxide (CO<sub>2</sub>) concentrations generally increase plant growth and C input to soil, suggesting that soil might help mitigate atmospheric CO<sub>2</sub> rise and global warming. But to what extent mitigation will occur is unclear. The large size of the soil C pool not only makes it a potential buffer against rising atmospheric CO<sub>2</sub>, but also makes it difficult to measure changes amid the existing background. Meta-analysis is one tool that can overcome the limited power of single studies. Four recent meta-analyses addressed this issue but reached somewhat different conclusions about the effect of elevated CO<sub>2</sub> on soil C accumulation, especially regarding the role of nitrogen (N) inputs. Here, we assess the extent of differences between these conclusions and propose a new analysis of the data. The four meta-analyses included different studies, derived different effect size estimates from common studies, used different weighting functions and metrics of effect size, and used different approaches to address nonindependence of effect sizes. Although all factors influenced the mean effect size estimates and subsequent inferences, the approach to independence had the largest influence. We recommend that meta-analysts critically assess and report choices about effect size metrics and weighting functions, and criteria for study selection and independence. Such decisions need to be justified carefully because they affect the basis for inference. Our new analysis, with a combined data set, confirms that the effect of elevated CO<sub>2</sub> on net soil C accumulation increases with the addition of N fertilizers. Although the effect at low N inputs was not significant, statistical power to detect biogeochemically important effect sizes at low N is limited, even with meta-analysis, suggesting the continued need for long-term experiments.
Hurteau MD, Hungate BA, Koch GW (2009) Accounting for risk in valuing forest carbon offsets. Carbon Balance and Management 4(1): 1-5.
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<p class="Para">Forests can sequester carbon dioxide, thereby reducing atmospheric concentrations and slowing global warming. In the U.S., forest carbon stocks have increased as a result of regrowth following land abandonment and in-growth due to fire suppression, and they currently sequester approximately 10% of annual US emissions. This ecosystem service is recognized in greenhouse gas protocols and cap-and-trade mechanisms, yet forest carbon is valued equally regardless of forest type, an approach that fails to account for risk of carbon loss from disturbance.</p>
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<p class="Para">Here we show that incorporating wildfire risk reduces the value of forest carbon depending on the location and condition of the forest. There is a general trend of decreasing risk-scaled forest carbon value moving from the northern toward the southern continental U.S.</p>
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<p class="Para">Because disturbance is a major ecological factor influencing long-term carbon storage and is often sensitive to human management, carbon trading mechanisms should account for the reduction in value associated with disturbance risk.</p>
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Langley JA, McKinley DC, Wolf AA, Hungate BA, Drake BG, Megonigal JP (2009) Priming depletes soil carbon and releases nitrogen in a scrub-oak ecosystem exposed to elevated CO 2. Soil Biology and Biochemistry 41(1): 54-60.
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Read PublicationElevated atmospheric CO<sub>2</sub> tends to stimulate plant productivity, which could either stimulate or suppress the processing of soil carbon, thereby feeding back to atmospheric CO<sub>2</sub> concentrations. We employed an acid-hydrolysis-incubation method and a net nitrogen-mineralization assay to assess stability of soil carbon pools and short-term nitrogen dynamics in a Florida scrub-oak ecosystem after six years of exposure to elevated CO<sub>2</sub>. We found that soil carbon concentration in the slow pool was 27% lower in elevated than ambient CO<sub>2</sub> plots at 0–10 cm depth. The difference in carbon mass was equivalent to roughly one-third of the increase in plant biomass that occurred in the same experiment. These results concur with previous reports from this ecosystem that elevated CO<sub>2</sub> stimulates microbial degradation of relatively stable soil organic carbon pools. Accordingly, elevated CO<sub>2</sub> increased net N mineralization in the 10–30 cm depth, which may increase N availability, thereby allowing for continued stimulation of plant productivity by elevated CO<sub>2</sub>. Our findings suggest that soil texture and climate may explain the differential response of soil carbon among various long-term, field-based CO<sub>2</sub> studies. Increased mineralization of stable soil organic carbon by a CO<sub>2</sub>-induced priming effect may diminish the terrestrial carbon sink globally.
Marks JC, Haden GA, Harrop BL, Reese EG, Keams JL, Watwood ME, Whitham TG (2009) Genetic and environmental controls of microbial communities on leaf litter in streams. Freshwater Biology 54(12): 2616-2627.
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1. Despite the importance of microorganisms for leaf litter decomposition in streams, little is known about which factors affect community composition of bacterial and fungal communities. Standard morphological techniques probably underestimate microbial diversity.
2. We used terminal restriction fragment length polymorphisms of the ITS regions for fungi, and the 16S region for bacteria, to compare fungal and bacterial communities on four cross types of cottonwood leaves (Populus fremontii, P. angustifolia, and their naturally occurring F1 and backcross hybrids). Decomposing leaves were studied in two Arizona rivers that differ in water chemistry and macroinvertebrates.
3. Hybridising cottonwoods are an ideal model system to test how genetic differences in leaf litter chemistry affect microbial communities because cross types have different decomposition rates and leaf litter chemistry. Leaves were incubated in litter bags for
2 weeks and brought to the laboratory for genetic analysis. Communities were analysed using non-metric multi dimensional scaling (NMDS) and diversity indices.
4. Fungal and bacterial communities differed between the two rivers, even when growing on identical substrates. There were also significant differences in microbial communities among the four cross types, indicating that genetically based differences in leaf litter translate to differences in microbial communities.
5. Diversity increased along the hybridising complex from P. fremontii to P. angustifolia, with hybrids showing intermediate values. Fungal and bacterial diversity were significantly higher on cross types with higher tannin concentrations and slower decomposition rates. 6. Environmental conditions most strongly structured microbial communities, but within an environment, genetic-based differences in leaf litter quality yielded differences in diversity and community structure.
7. Molecular tools are making it possible to understand patterns of microbial diversity in river ecosystems, paving the way for a better understanding of how differences in microbial species affect ecosystem processes and higher trophic levels.
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McKinley DC, Romero JC, Hungate BA, Drake BG, Megonigal JP (2009) Does deep soil N availability sustain long‐term ecosystem responses to elevated CO2?. Global Change Biology 15(5): 2035-2048.
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Read PublicationA scrub-oak woodland has maintained higher aboveground biomass accumulation after 11 years of atmospheric CO<sub>2</sub> enrichment (ambient +350 μmol CO<sub>2</sub> mol<sup>−1</sup>), despite the expectation of strong nitrogen (N) limitation at the site. We hypothesized that changes in plant available N and exploitation of deep sources of inorganic N in soils have sustained greater growth at elevated CO<sub>2</sub>. We employed a suite of assays performed in the sixth and 11th year of a CO<sub>2</sub> enrichment experiment designed to assess soil N dynamics and N availability in the entire soil profile. In the 11th year, we found no differences in gross N flux, but significantly greater microbial respiration (<em>P≤</em>0.01) at elevated CO<sub>2</sub>. Elevated CO<sub>2</sub> lowered extractable inorganic N concentrations (<em>P</em>=0.096) considering the whole soil profile (0–190 cm). Conversely, potential net N mineralization, although not significant in considering the entire profile (<em>P</em>=0.460), tended to be greater at elevated CO<sub>2</sub>. Ion-exchange resins placed in the soil profile for approximately 1 year revealed that potential N availability at the water table was almost 3 × greater than found elsewhere in the profile, and we found direct evidence using a <sup>15</sup>N tracer study that plants took up N from the water table. Increased microbial respiration and shorter mean residence times of inorganic N at shallower depths suggests that enhanced SOM decomposition may promote a sustained supply of inorganic N at elevated CO<sub>2</sub>. Deep soil N availability at the water table is considerable, and provides a readily available source of N for plant uptake. Increased plant growth at elevated CO<sub>2</sub> in this ecosystem may be sustained through greater inorganic N supply from shallow soils and N uptake from deep soil.
Meulbauer JD, LeRoy CJ, Lovett JM, Flaccus KK, Vlieg JK, Marks JC (2009) Short-term responses of decomposers to flow restoration in Fossil Creek, Arizona, USA. Hydrobiologia 618(1): 35-45.
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Read PublicationDam decommissioning projects, although numerous, rarely include complete sets of data before and after restoration for evaluating the ecological consequences of such projects. In this study, we used a before-after control-impact (BACI) design to assess changes in leaf litter decomposition and associated macroinvertebrate and fungal decomposers following dam decommissioning in Fossil Creek, Arizona, USA. Leaf litterbags were deployed in a relatively pristine site above the dam and a highly disturbed site below the dam where over 95% of the flow was previously diverted for hydropower generation. Leaf litter decomposition was significantly slower below the dam both measurement years (pre- and post-restoration) with no site-year interaction, indicating that decomposition in this stream section was not affected by increased flow. In contrast, both macroinvertebrates and fungi differed significantly above and below the dam prior to restoration but were similar post-restoration, supporting the concept that decomposer communities can quickly rebound following reintroduction of full flow. Our results indicate that some aquatic ecosystem variables can return to a more natural state following ecological restoration activities such as water flow restoration.
Montes-Helu MC, Kolb T, Dore S, Sullivan B, Hart SC, Koch G, Hungate BA (2009) Persistent effects of fire-induced vegetation change on energy partitioning and evapotranspiration in ponderosa pine forests. Agricultural and Forest Meteorology 149(3): 491-500.
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Read PublicationWe compared energy fluxes between a site converted from ponderosa pine (<em>Pinus ponderosa</em>) forest to sparse grassland by a severe wildfire 10 years ago and a nearby, unburned forest. We used eddy covariance and associated instruments to measure total radiation, net radiation, albedo, and fluxes of energy into latent heat, sensible heat, and the soil. Total radiation, vapor pressure deficit, and air temperature were similar for each site. Compared to the unburned site, net radiation efficiency (net radiation/total radiation) was 30% lower and albedo 30% higher at the burned site. The magnitude of sensible and latent heats varied seasonally at both sites. Sensible heat was the major component of the energy balance in cold or dry seasons, whereas latent heat was the major component in the warm and wet season. Soil heat flux was the smallest in magnitude of the measured energy fluxes. Compared with the unburned forest, the burn-created grassland generally had lower sensible and latent heats, but greater soil heat flux for both soil cooling in winter and warming in summer. The grassland had similar maximum air temperature as the forest, and warmer surface soil temperature during the summer. Thus, the lower albedo and greater sensible heat of the forest did not produce a warmer site compared with the grassland, apparently because of the cooling effect of greater latent heat in the forest. Our results suggest only small changes in site air temperature, but larger changes in site surface soil temperature by shifts from forest to grassland caused by severe fire in northern Arizona ponderosa pine forests.
Osterkamp TE, Jorgenson MT, Schuur EAG, Shur YL, Kanevskiy MZ, Vogel JG, Tumskoy VE (2009) Physical and ecological changes associated with warming permafrost and thermokarst in interior Alaska. Permafrost and Periglacial Processes 20(3): 235-256.
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Read PublicationObservations and measurements were made of physical and ecological changes that have occurred since 1985 at a tundra site near Healy, Alaska. Air temperatures decreased (1985 through 1999) while permafrost warmed and thawed creating thermokarst terrain, probably as a result of increased snow depths. Permafrost, active layer and ground-ice conditions at the Healy site are the result of the interaction of climatic, ecologic and other factors. The slow accumulation of ground ice in an intermediate permafrost layer formed by upward freezing from the permafrost surface leads to long-term differential frost heave and microrelief. When ground ice in the permafrost melts, the ground surface settles differentially resulting in thermokarst terrain (pits, gullies). Windblown snow fills the thermokarst depressions causing further warming and thawing of the underlying permafrost — a positive feedback effect that enhances permafrost degradation. Thermokarst-induced changes in relief alter the near-surface hydrology and ecological processes. Changes in vegetation included differential tussock growth and mortality and a shift in moss species abundance and relative productivity, depending on microtopographic position created by the thermokarst terrain. Water redistribution towards thermokarst depressions caused adjacent higher areas to become drier and resulted in increased moss mortality and shrub abundance.
Seiler TJ, Rasse DP, Li J, Dijkstra P, Anderson HP, Johnson DP, Powell TL, Hungate BA, Hinkle C, Drake BG (2009) Disturbance, rainfall and contrasting species responses mediated aboveground biomass response to 11 years of CO2 enrichment in a Florida scrub‐oak ecosystem. Global Change Biology 15(2): 356-367.
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Read PublicationThis study reports the aboveground biomass response of a fire-regenerated Florida scrub-oak ecosystem exposed to elevated CO<sub>2</sub> (1996–2007), from emergence after fire through canopy closure. Eleven years exposure to elevated CO<sub>2</sub> caused a 67% increase in aboveground shoot biomass. Growth stimulation was sustained throughout the experiment; although there was significant variability between years. The absolute stimulation of aboveground biomass generally declined over time, reflecting increasing environmental limitations to long-term growth response. Extensive defoliation caused by hurricanes in September 2004 was followed by a strong increase in shoot density in 2005 that may have resulted from reopening the canopy and relocating nitrogen from leaves to the nutrient-poor soil. Biomass response to elevated CO<sub>2</sub> was driven primarily by stimulation of growth of the dominant species, <em>Quercus myrtifolia</em>, while <em>Quercus geminata</em>, the other co-dominant oak, displayed no significant CO<sub>2</sub> response. Aboveground growth also displayed interannual variation, which was correlated with total annual rainfall. The rainfall × CO<sub>2</sub> interaction was partially masked at the community level by species-specific responses: elevated CO<sub>2</sub> had an ameliorating effect on <em>Q. myrtifolia</em> growth under water stress. The results of this long-term study not only show that atmospheric CO<sub>2</sub> concentration had a consistent stimulating effect on aboveground biomass production, but also showed that available water is the primary driver of interannual variation in shoot growth and that the long-term response to elevated CO<sub>2</sub> may have been caused by other factors such as nutrient limitation and disturbance.
Stiling P, Moon D, Rossi A, Hungate BA, Drake B (2009) Seeing the forest for the trees: long‐term exposure to elevated CO2 increases some herbivore densities. Global Change Biology 15(8): 1895-1902.
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Read PublicationThe effects of elevated CO<sub>2</sub> on plant growth and insect herbivory have been frequently investigated over the past 20 years. Most studies have shown an increase in plant growth, a decrease in plant nitrogen concentration, an increase in plant secondary metabolites and a decrease in herbivory. However, such studies have generally overlooked the fact that increases in plant production could cause increases of herbivores per unit area of habitat. Our study investigated leaf production, herbivory levels and herbivore abundance per unit area of leaf litter in a scrub-oak system at Kennedy Space Center, Florida, under conditions of ambient and elevated CO<sub>2</sub>, over an 11-year period, from 1996 to 2007. In every year, herbivory, that is leafminer and leaftier abundance per 200 leaves, was lower under elevated CO<sub>2</sub> than ambient CO<sub>2</sub> for each of three species of oaks, <em>Quercus myrtifolia, Quercus chapmanii</em> and <em>Quercus geminata</em>. However, leaf litter production per 0.1143 m<sup>2</sup> was greater under elevated CO<sub>2</sub> than ambient CO<sub>2</sub> for <em>Q. myrtifolia</em> and <em>Q. chapmanii</em>, and this difference increased over the 11 years of the study. Leaf production of <em>Q. geminata</em> under elevated CO<sub>2</sub> did not increase. Leafminer densities per 0.1143 m<sup>2</sup> of litterfall for <em>Q. myrtifolia</em> and <em>Q. chapmanii</em> were initially lower under elevated CO<sub>2</sub>. However, shortly after canopy closure in 2001, leafminer densities per 0.1143 m<sup>2</sup> of litter fall became higher under elevated CO<sub>2</sub> and remained higher for the remainder of the experiment. Leaftier densities per 0.1143 m<sup>2</sup> were also higher under elevated CO<sub>2</sub> for <em>Q. myrtifolia</em> and <em>Q. chapmanii</em> over the last 6 years of the experiment. There were no differences in leafminer or leaftier densities per 0.1143 m<sup>2</sup> of litter for <em>Q. geminata</em>. These results show three phenomena. First, they show that elevated CO<sub>2</sub> decreases herbivory on all oak species in the Florida scrub-oak system. Second, despite lower numbers of herbivores per 200 leaves in elevated CO<sub>2</sub>, increased leaf production resulted in higher herbivore densities per unit area of leaf litter for two oak species. Third, they corroborate other studies which suggest that the effects of elevated CO<sub>2</sub> on herbivores are species specific, meaning they depend on the particular plant species involved. Two oak species showed increases in leaf production and herbivore densities per 0.1143 m<sup>2</sup> in elevated CO<sub>2</sub> over time while another oak species did not. Our results point to a future world of elevated CO<sub>2</sub> where, despite lower plant herbivory, some insect herbivores may become more common.
Vogel J, Schuur EAG, Trucco C, Lee H (2009) Response of CO2 exchange in a tussock tundra ecosystem to permafrost thaw and thermokarst development. Journal of Geophysical Research: Biogeosciences 114: G4.
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Read PublicationClimate change in high latitudes can lead to permafrost thaw, which in ice-rich soils can result in ground subsidence, or thermokarst. In interior Alaska, we examined seasonal and annual ecosystem CO<sub>2</sub> exchange using static and automatic chamber measurements in three areas of a moist acidic tundra ecosystem undergoing varying degrees of permafrost thaw and thermokarst development. One site had extensive thermokarst features, and historic aerial photography indicated it was present at least 50 years prior to this study. A second site had a moderate number of thermokarst features that were known to have developed concurrently with permafrost warming that occurred 15 years prior to this study. A third site had a minimal amount of thermokarst development. The areal extent of thermokarst features reflected the seasonal thaw depth. The “extensive” site had the deepest seasonal thaw depth, and the “moderate” site had thaw depths slightly, but not significantly deeper than the site with “minimal” thermokarst development. Greater permafrost thaw corresponded to significantly greater gross primary productivity (GPP) at the moderate and extensive thaw sites as compared to the minimal thaw site. However, greater ecosystem respiration (R<sub>eco</sub>) during the spring, fall, and winter resulted in the extensive thaw site being a significant net source of CO<sub>2</sub> to the atmosphere over 3 years, while the moderate thaw site was a CO<sub>2</sub> sink. The minimal thaw site was near CO<sub>2</sub> neutral and not significantly different from the extensive thaw site. Thus after permafrost thaw, initial periods of increased GPP and net CO<sub>2</sub> uptake could be offset by elevated R<sub>eco</sub>during the winter, spring, and fall.
2008
Dijkstra P, LaViolette CM, Coyle JS, Doucett RR, Schwartz E, Hart SC, Hungate BA (2008) 15N enrichment as an integrator of the effects of C and N on microbial metabolism and ecosystem function. Ecology Letters 11(4): 389-397.
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Read PublicationOrganic carbon (C) and nitrogen (N) are essential for heterotrophic soil microorganisms, and their bioavailability strongly influences ecosystem C and N cycling. We show here that the natural <sup>15</sup>N abundance of the soil microbial biomass is affected by both the availability of C and N and ecosystem N processing. Microbial <sup>15</sup>N enrichment correlated negatively with the C : N ratio of the soil soluble fraction and positively with net N mineralization for ecosystems spanning semiarid, temperate and tropical climates, grassland and forests, and over four million years of ecosystem development. In addition, during soil incubation, large increases in microbial <sup>15</sup>N enrichment corresponded to high net N mineralization rates. These results support the idea that the N isotope composition of an organism is determined by the balance between N assimilation and dissimilation. Thus, <sup>15</sup>N enrichment of the soil microbial biomass integrates the effects of C and N availability on microbial metabolism and ecosystem processes.
Dore S, Kolb TE, Montes‐Helu M, Sullivan BW, Winslow WD, Hart SC, Kaye JP, Koch GW, Hungate BA (2008) Long‐term impact of a stand‐replacing fire on ecosystem CO2 exchange of a Ponderosa pine forest. Global Change Biology 14(8): 1801-1820.
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Read PublicationPonderosa pine (<em>Pinus ponderosa</em>) forests of the southwestern United States are a mosaic of stands where undisturbed forests are carbon sinks, and stands recovering from wildfires may be sources of carbon to the atmosphere for decades after the fire. However, the relative magnitude of these sinks and sources has never been directly measured in this region, limiting our understanding of the role of fire in regional and US carbon budgets. We used the eddy covariance technique to measure the CO<sub>2</sub> exchange of two forest sites, one burned by fire in 1996, and an unburned forest. The fire was a high-intensity stand-replacing burn that killed all trees. Ten years after the fire, the burned site was still a source of CO<sub>2</sub> to the atmosphere [109±6 (SEM) g C m<sup>−2</sup> yr<sup>−1</sup>], whereas the unburned site was a sink (−164±23 g C m<sup>−2</sup> yr<sup>−1</sup>). The fire reduced total carbon storage and shifted ecosystem carbon allocation from the forest floor and living biomass to necromass. Annual ecosystem respiration was lower at the burned site (480±5 g C m<sup>−2</sup> yr<sup>−1</sup>) than at the unburned site (710±54 g C m<sup>−2</sup> yr<sup>−1</sup>), but the difference in gross primary production was even larger (372±13 g C m<sup>−2</sup> yr<sup>−1</sup> at the burned site and 858±37 g C m<sup>−2</sup> yr<sup>−1</sup>at the unburned site). Water availability controlled carbon flux in the warm season at both sites, and the burned site was a source of carbon in all months, even during the summer, when wet and warm conditions favored respiration more than photosynthesis. Our study shows that carbon losses following stand-replacing fires in ponderosa pine forests can persist for decades due to slow recovery of the gross primary production. Because fire exclusion is becoming increasingly difficult in dry western forests, a large US forest carbon sink could shift to a decadal-scale carbon source.
Duval BD, Hungate BA (2008) Soil science: Scavenging for scrap metal. Nature Geoscience 1(4): 213-214.
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Read Publication<div> Organisms acquire some elements from the environment with ease. Diffusion alone often provides enough carbon dioxide, oxygen and water. But getting other elements requires more effort, spurring unique evolutionary adaptations: instead of taking up nutrients from the soil, some plants in acidic bogs trap insects to obtain nitrogen and phosphorus 1; geophagy — or eating dirt — may sometimes be important for acquiring iron by primates 2; plants and microorganisms secrete compounds that liberate phosphorus from unavailable forms in soil 3; and many bacteria secrete metal-scavenging compounds called siderophores to capture iron and copper 4,5. Evidence has mounted that molybdenum is also specifically targeted 6. On page 243 of this issue, Bellenger and colleagues 7 confirm this, showing that siderophores produced by the nitrogen-fixing bacterium Azotobacter vinelandiibind with molybdenum and vanadium in the laboratory, promoting uptake of these metals.</div>
Hurteau MD, Koch GW, Hungate BA (2008) Carbon protection and fire risk reduction: toward a full accounting of forest carbon offsets. Frontiers in Ecology and the Environment 6(9): 493-498.
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Read PublicationManagement of forests for carbon uptake is an important tool in the effort to slow the increase in atmospheric CO<sub>2</sub> and global warming. However, some current policies governing forest carbon credits actually promote avoidable CO<sub>2</sub> release and punish actions that would increase long-term carbon storage. In fire-prone forests, management that reduces the risk of catastrophic carbon release resulting from stand-replacing wild-fire is considered to be a CO<sub>2</sub> source, according to current accounting practices, even though such management may actually increase long-term carbon storage. Examining four of the largest wildfires in the US in 2002, we found that, for forest land that experienced catastrophic stand-replacing fire, prior thinning would have reduced CO<sub>2</sub> release from live tree biomass by as much as 98%. Altering carbon accounting practices for forests that have historically experienced frequent, low-severity fire could provide an incentive for forest managers to reduce the risk of catastrophic fire and associated large carbon release events.
Menyailo OV, Hungate BA, Abraham W-R, Conrad R (2008) Changing land use reduces soil CH4 uptake by altering biomass and activity but not composition of high‐affinity methanotrophs. Global Change Biology 14(10): 2405-2419.
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Read PublicationForest ecosystems assimilate more CO<sub>2</sub> from the atmosphere and store more carbon in woody biomass than most nonforest ecosystems, indicating strong potential for afforestation to serve as a carbon management tool. However, converting grasslands to forests could affect ecosystem–atmosphere exchanges of other greenhouse gases, such as nitrous oxide and methane (CH<sub>4</sub>), effects that are rarely considered. Here, we show that afforestation on a well-aerated grassland in Siberia reduces soil CH<sub>4</sub> uptake by a factor of 3 after 35 years of tree growth. The decline in CH<sub>4</sub> oxidation was observed both in the field and in laboratory incubation studies under controlled environmental conditions, suggesting that not only physical but also biological factors are responsible for the observed effect. Using incubation experiments with <sup>13</sup>CH<sub>4</sub> and tracking <sup>13</sup>C incorporation into bacterial phospholipid fatty acid (PLFA), we found that, at low CH<sub>4</sub> concentrations, most of the <sup>13</sup>C was incorporated into only two PLFAs, 18 : 1ω7 and 16 : 0. High CH<sub>4</sub> concentration increased total <sup>13</sup>C incorporation and the number of PLFA peaks that became labeled, suggesting that the microbial assemblage oxidizing CH<sub>4</sub> shifts with ambient CH<sub>4</sub> concentration. Forests and grasslands exhibited similar labeling profiles for the high-affinity methanotrophs, suggesting that largely the same general groups of methanotrophs were active in both ecosystems. Both PLFA concentration and labeling patterns indicate a threefold decline in the biomass of active methanotrophs due to afforestation, but little change in the methanotroph community. Because the grassland consumed CH<sub>4</sub> at a rate five times higher than forest soils under laboratory conditions, we concluded that not only biomass but also cell-specific activity was higher in grassland than in afforested plots. While the decline in biomass of active methanotrophs can be explained by site preparation (plowing), inorganic N (especially NH<sub>4</sub><sup>+</sup>) could be responsible for the change in cell-specific activity. Overall, the negative effect of afforestation of upland grassland on soil CH<sub>4</sub> uptake can be largely explained by the reduction in biomass and to a lesser extent by reduced cell-specific activity of CH<sub>4</sub>-oxidizing bacteria.
Selmants PC, Hart SC, Boyle SI, Gehring CA, Hungate BA (2008) Restoration of a ponderosa pine forest increases soil CO2 efflux more than either water or nitrogen additions. Journal of Applied Ecology 45(3): 913-920.
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Read Publication<div data-canvas-width="658.8216666666666">1. Ecological restoration often involves returning ecosystem structure to some predisturbance</div>
<div data-canvas-width="679.9833333333332">reference state, but ecosystem function must also recover if restoration efforts are to be self-sustaining</div>
<div data-canvas-width="680.0049999999998">over the long term. In the south-western United States, ponderosa pine forest structure was altered</div>
<div data-canvas-width="679.9983333333332">by disruption of the fire regime following Euro-American settlement. Forest structure is now being</div>
<div data-canvas-width="679.9983333333331">restored to presettlement conditions through the application of thinning and burning treatments.</div>
<div data-canvas-width="663.6866666666666">However, the effects of these treatments on below-ground ecosystem processes remain unclear.</div>
<div data-canvas-width="658.8149999999998">2. We conducted a water and nitrogen (N) addition experiment in adjacent restored and unrestored</div>
<div data-canvas-width="366.61499999999995">ponderosa pine stands and compared soil CO2 efflux in response to these treatments over a 13-month</div>
<div data-canvas-width="680.0016666666666">period. Our goals were to (i) quantify water and N limitation to below-ground carbon (C) cycling</div>
<div data-canvas-width="680.0066666666665">in contemporary high-density ponderosa pine forests; and (ii) determine if restoration alleviates</div>
<div data-canvas-width="170.73000000000002">water and N limitations.</div>
<div data-canvas-width="158.11333333333332">3. Restoration thinning and burning increased soil CO2 efflux, along with surface soil watercontent, temperature and herbaceous fine root biomass, while total fine root biomass decreased asa result of restoration.</div>
<div data-canvas-width="658.8283333333333">4. Water and N additions increased C flux from soils to a similar degree in both restored and</div>
<div data-canvas-width="680.0166666666668">unrestored ponderosa pine stands, but the increase was relatively small when compared to that</div>
<div data-canvas-width="180.79666666666665">stimulated by restoration.</div>
<div data-canvas-width="476.5566666666667">5. Synthesis and applications. An understanding of how ecosystem processes respond to treatments</div>
<div data-canvas-width="680.0016666666667">designed to restore ecosystem structure is critical in ensuring the long-term success of restoration</div>
<div data-canvas-width="680.0033333333333">efforts. Here we show that, although water and N stimulate C flux from soils in these semi-arid</div>
<div data-canvas-width="679.9999999999999">forests, restoration treatments have a much greater effect on soil C balance than increased water and N availability by themselves. This suggests that increased quality of C inputs from a recovering</div>
<div data-canvas-width="680.17">understorey herbaceous community is a key component of restoring ecosystem function (e.g.</div>
<div data-canvas-width="460.53999999999996">below-ground C cycling) in south-western ponderosa pine forests.</div>
2007
Carney KM, Hungate BA, Drake BG, Megonigal JP (2007) Altered soil microbial community at elevated CO2 leads to loss of soil carbon. Proceedings of the National Academy of Sciences 104(12): 4990-4995.
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Read PublicationIncreased carbon storage in ecosystems due to elevated CO<sub>2</sub> may help stabilize atmospheric CO<sub>2</sub> concentrations and slow global warming. Many field studies have found that elevated CO<sub>2</sub> leads to higher carbon assimilation by plants, and others suggest that this can lead to higher carbon storage in soils, the largest and most stable terrestrial carbon pool. Here we show that 6 years of experimental CO<sub>2</sub> doubling reduced soil carbon in a scrub-oak ecosystem despite higher plant growth, offsetting ≈52% of the additional carbon that had accumulated at elevated CO<sub>2</sub> in aboveground and coarse root biomass. The decline in soil carbon was driven by changes in soil microbial composition and activity. Soils exposed to elevated CO<sub>2</sub> had higher relative abundances of fungi and higher activities of a soil carbon-degrading enzyme, which led to more rapid rates of soil organic matter degradation than soils exposed to ambient CO<sub>2</sub>. The isotopic composition of microbial fatty acids confirmed that elevated CO<sub>2</sub> increased microbial utilization of soil organic matter. These results show how elevated CO<sub>2</sub>, by altering soil microbial communities, can cause a potential carbon sink to become a carbon source.
Carter CD, Marks JC (2007) Influences of travertine dam formation on leaf litter decomposition and algal accrual. Hydrobiologia 575(1): 329-341.
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Read PublicationAt the time of this study Fossil Creek was being considered as a site for the restoration of a native fish assemblage, however there was concern amongst fisheries managers about the stream being food limited due to calcium carbonate (travertine) deposition. To evaluate the effects of travertine deposition on the aquatic food base we used leaf litterbags to compare decomposition rates and nutrient diffusing artificial substrates to compare algal accrual rates and nutrient limitation between two distinct reaches in Fossil creek: a travertine dam forming reach and a reach without travertine dam formation (riffle-pool reach). Decomposition was significantly faster in the travertine dam forming reach than in the riffle-pool reach. Macroinvertebrates in the leaf packs were more diverse in the travertine reach but more abundant in the riffle-pool reach. Algae accrued more quickly in the travertine reach than in the riffle-pool reach and only responded to nutrient enrichment in the travertine reach. This study was conducted prior to a hydroelectric dam decommissioning project in Fossil Creek where full flows were reintroduced back into the stream after a century of diversion. Our results suggest concurrent increases in algal productivity, decomposition, and macroinvertebrate diversity in the next decade as travertine dams rebuild, providing a richer food base for fish and other aquatic organisms.
Dinger EC, Marks JC (2007) Effects of high levels of antimycin A on aquatic invertebrates in a warmwater Arizona Stream. North American Journal of Fisheries Management 27(4): 1243-1256.
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Read PublicationRestoration of native fish to freshwater habitats often requires nonnative fish removal via chemicals such as antimycin A. Despite widespread use, there are limited field studies quantifying the effects of antimycin A on aquatic macroinvertebrates. We studied the immediate and short-term effects of antimycin A on macroinvertebrates during a fish renovation project in Fossil Creek, Arizona. We employed before–after control–impact (BACI) designs to measure the effects of antimycin A (at extraordinarily high levels of >54 and >100 μg/L) on macroinvertebrate drift, density, and species composition. We used the Hilsenhoff biotic index, a measure of invertebrate pollution tolerance, to study changes in species composition. At the highest dose (>100 μg/L), drift was five times the pretreatment drift level and invertebrate standing stocks in pools and riffles decreased immediately. Densities rebounded in riffles within 5 months but remained depressed in pools. At the lower concentration (>54 μg/L), macroinvertebrate mortality, measured as increased drift, was 24 times the pretreatment level. At this lower concentration, however, macroinvertebrate densities in the benthos were not reduced. Under both concentrations, species composition shifted toward more tolerant species. Although antimycin A effects were mostly short term, several species were locally extirpated. We found no explanation for the loss of some species over others. These results indicate that there is a high end concentration at which antimycin A can have deleterious effects on aquatic invertebrates. We caution managers contemplating the use of antimycin A in fish restoration to consider the risks to macroinvertebrates. We suggest the use of pretreatment surveys and bioassays at anticipated treatment levels to predict the effects upon macroinvertebrates, especially sensitive species. Where there are sensitive species, steps should be taken to mitigate effects.
Doucette RR, Marks JC, Blinn DW, Caron M, Hungate BA (2007) Measuring terrestrial subsidies to aquatic food webs using stable isotopes of hydrogen. Ecology 88(6):1587-1592.
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Read PublicationUnderstanding river food webs requires distinguishing energy derived from primary production in the river itself (autochthonous) from that produced externally (allochthonous), yet there are no universally applicable and reliable techniques for doing so. We compared the natural abundance stable isotope ratios of hydrogen (δD) of allochthonous and autochthonous energy sources in four different aquatic ecosystems. We found that autochthonous organic matter is uniformly far more depleted in deuterium (lower δD values) than allochthonous: an average difference of ∼100‰. We also found that organisms at higher trophic levels, including both aquatic invertebrates and fish, have δD values intermediate between aquatic algae and terrestrial plants. The consistent differences between leaves and algae in δD among these four watersheds, along with the intermediate values in higher trophic levels, indicate that natural abundance hydrogen isotope signatures are a powerful tool for partitioning energy flow in aquatic ecosystems.
Hungate BA, Hart SC, Selmants PC, Boyle SI, Gehring CA (2007) Soil responses to management, increased precipitation, and added nitrogen in ponderosa pine forests. Ecological Applications 17(5): 1352-1365.
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Read PublicationForest management, climatic change, and atmospheric N deposition can affect soil biogeochemistry, but their combined effects are not well understood. We examined the effects of water and N amendments and forest thinning and burning on soil N pools and fluxes in ponderosa pine forests near Flagstaff, Arizona (USA). Using a <sup>15</sup>N-depleted fertilizer, we also documented the distribution of added N into soil N pools. Because thinning and burning can increase soil water content and N availability, we hypothesized that these changes would alleviate water and N limitation of soil processes, causing smaller responses to added N and water in the restored stand. We found little support for this hypothesis. Responses of fine root biomass, potential net N mineralization, and the soil microbial N to water and N amendments were mostly unaffected by stand management. Most of the soil processes we examined were limited by N and water, and the increased N and soil water availability caused by forest restoration was insufficient to alleviate these limitations. For example, N addition caused a larger increase in potential net nitrification in the restored stand, and at a given level of soil N availability, N addition had a larger effect on soil microbial N in the restored stand. Possibly, forest restoration increased the availability of some other limiting resource, amplifying responses to added N and water. Tracer N recoveries in roots and in the forest floor were lower in the restored stand. Natural abundance δ<sup>15</sup>N of labile soil N pools were higher in the restored stand, consistent with a more open N cycle. We conclude that thinning and burning open up the N cycle, at least in the short term, and that these changes are amplified by enhanced precipitation and N additions. Our results suggest that thinning and burning in ponderosa pine forests will not increase their resistance to changes in soil N dynamics resulting from increased atmospheric N deposition or increased precipitation due to climatic change. Restoration plans should consider the potential impact on long-term forest productivity of greater N losses from a more open N cycle, especially during the period immediately after thinning and burning.
Li J, Powell TL, Seiler TJ, Johnson DP, Anderson HP, Bracho R, Hungate BA, Hinkle CR, Drake BG (2007) Impacts of hurricane Frances on Florida scrub‐oak ecosystem processes: Defoliation, net CO2 exchange and interactions with elevated CO2. Global Change Biology 13(6): 1101-1113.
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Read PublicationHurricane disturbances have profound impacts on ecosystem structure and function, yet their effects on ecosystem CO<sub>2</sub> exchange have not been reported. In September 2004, our research site on a fire-regenerated scrub-oak ecosystem in central Florida was struck by Hurricane Frances with sustained winds of 113 km h<sup>−1</sup> and wind gusts as high as 152 km h<sup>−1</sup>. We quantified the hurricane damage on this ecosystem resulting from defoliation: we measured net ecosystem CO<sub>2</sub> exchange, the damage and recovery of leaf area, and determined whether growth in elevated carbon dioxide concentration in the atmosphere (<em>C</em><sub>a</sub>) altered this disturbance. The hurricane decreased leaf area index (LAI) by 21%, which was equal to 60% of seasonal variation in canopy growth during the previous 3 years, but stem damage was negligible. The reduction in LAI led to a 22% decline in gross primary production (GPP) and a 25% decline in ecosystem respiration (<em>R</em><sub>e</sub>). The compensatory declines in GPP and <em>R</em><sub>e</sub> resulted in no significant change in net ecosystem production (NEP). Refoliation began within a month after the hurricane, although this period was out of phase with the regular foliation period, and recovered 20% of the defoliation loss within 2.5 months. Full recovery of LAI, ecosystem CO<sub>2</sub> assimilation, and ecosystem respiration did not occur until the next growing season. Plants exposed to elevated <em>C</em><sub>a</sub> did not sustain greater damage, nor did they recover faster than plants grown under ambient <em>C</em><sub>a</sub>. Thus, our results indicate that hurricanes capable of causing significant defoliation with negligible damage to stems have negligible effects on NEP under current or future CO<sub>2</sub>-enriched environment.
Li JH, Johnson DP, Dijkstra P, Hungate BA, Hinkle CR, Drake BG (2007) Elevated CO2 mitigates the adverse effects of drought on daytime net ecosystem CO2 exchange and photosynthesis in a Florida scrub-oak ecosystem. Photosynthetica 45(1): 51-58.
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Read PublicationDrought is a normal, recurrent feature of climate. In order to understand the potential effect of increasing atmospheric CO<sub>2</sub> concentration (<em class="EmphasisTypeItalic ">C</em> <sub>a</sub>) on ecosystems, it is essential to determine the combined effects of drought and elevated <em class="EmphasisTypeItalic ">C</em> <sub>a</sub> (EC) under field conditions. A severe drought occurred in Central Florida in 1998 when precipitation was 88 % less than the average between 1984 and 2002. We determined daytime net ecosystem CO<sub>2</sub> exchange (NEE) before, during, and after the drought in the Florida scrub-oak ecosystem exposed to doubled <em class="EmphasisTypeItalic ">C</em> <sub>a</sub> in open-top chamber since May 1996. We measured diurnal leaf net photosynthetic rate (<em class="EmphasisTypeItalic ">P</em> <sub>N</sub>) of <em class="EmphasisTypeItalic ">Quercus myrtifolia</em> Willd, the dominant species, during and after the drought. Drought caused a midday depression in NEE and <em class="EmphasisTypeItalic ">P</em> <sub>N</sub> at ambient CO<sub>2</sub> concentration (AC) and EC. EC mitigated the midday depression in NEE by about 60 % compared to AC and the effect of EC on leaf <em class="EmphasisTypeItalic ">P</em> <sub>N</sub> was similar to its effect on NEE. Growth in EC lowered the sensitivity of NEE to air vapor pressure deficit under drought. Thus EC would help the scrub-oak ecosystem to survive the consequences of the effects of rising atmospheric CO<sub>2</sub> on climate change, including increased frequency of drought, while simultaneously sequestering more anthropogenic carbon.
Marks, JC (2007) Down Go the Dams. Scientific American 296(3): 66-71.
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Read PublicationThe article focuses on the decommissioning of dams across the United States. Many dams were built after World War II to provide sources of hydraulic power. Now that these power suppliers make up little of the nation's power resources, policy makers, conservationists, and environmentalists are opting to decommission the dams. Even though the natural ecology is expected to flourish from the decommissioning of dams, scientists are worried that there may be some negative effects.
Pagel Brown AL, Day FP, Hungate BA, Drake BG, Hinkle CR (2007) Root biomass and nutrient dynamics in a scrub-oak ecosystem under the influence of elevated atmospheric CO2. Plant and Soil 292(1-2): 219-232.
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Read PublicationElevated CO<sub>2</sub> can increase fine root biomass but responses of fine roots to exposure to increased CO<sub>2</sub> over many years are infrequently reported. We investigated the effect of elevated CO<sub>2</sub> on root biomass and N and P pools of a scrub-oak ecosystem on Merritt Island in Florida, USA, after 7 years of CO<sub>2</sub> treatment. Roots were removed from 1-m deep soil cores in 10-cm increments, sorted into different categories (<0.25 mm, 0.25–1 mm, 1–2 mm, 2 mm to 1 cm, >1 cm, dead roots, and organic matter), weighed, and analyzed for N, P and C concentrations. With the exception of surface roots <0.25 mm diameter, there was no effect of elevated CO<sub>2</sub> on root biomass. There was little effect on C, N, or P concentration or content with the exception of dead roots, and <0.25 mm and 1–2 mm diameter live roots at the surface. Thus, fine root mass and element content appear to be relatively insensitive to elevated CO<sub>2</sub>. In the top 10 cm of soil, biomass of roots with a diameter of <0.25 mm was depressed by elevated CO<sub>2</sub>. Elevated CO<sub>2</sub> tended to decrease the mass and N content of dead roots compared to ambient CO<sub>2</sub>. A decreased N concentration of roots <0.25 mm and 1–2 mm in diameter under elevated CO<sub>2 </sub>may indicate reduced N supply in the elevated CO<sub>2</sub> treatment. Our study indicated that elevated CO<sub>2</sub> does not increase fine root biomass or the pool of C in fine roots. In fact, elevated CO<sub>2</sub> tends to reduce biomass and C content of the most responsive root fraction (<0.25 mm roots), a finding that may have more general implications for understanding C input into the soil at higher atmospheric CO<sub>2</sub> concentrations.
Schwartz E, Blazewicz S, Doucett R, Hungate BA, Hart SC, Dijkstra P (2007) Natural abundance δ15N and δ13C of DNA extracted from soil. Soil Biology and Biochemistry 39(12): 3101-3107.
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Read PublicationWe report the first simultaneous measurements of <em>δ</em><sup>15</sup>N and <em>δ</em><sup>13</sup>C of DNA extracted from surface soils. The isotopic composition of DNA differed significantly among nine different soils. The <em>δ</em><sup>13</sup>C and <em>δ</em><sup>15</sup>N of DNA was correlated with <em>δ</em><sup>13</sup>C and <em>δ</em><sup>15</sup>N of soil, respectively, suggesting that the isotopic composition of DNA is strongly influenced by the isotopic composition of soil organic matter. However, in all samples DNA was enriched in <sup>13</sup>C relative to soil, indicating microorganisms fractionated C during assimilation or preferentially used <sup>13</sup>C enriched substrates. Enrichment of DNA in <sup>15</sup>N relative to soil was not consistently observed, but there were significant differences between <em>δ</em><sup>15</sup>N of DNA and <em>δ</em><sup>15</sup>N of soil for three different sites, suggesting microorganisms are fractionating N or preferentially using N substrates at different rates across these contrasting ecosystems. There was a strong linear correlation between <em>δ</em><sup>15</sup>N of DNA and <em>δ</em><sup>15</sup>N of the microbial biomass, which indicated DNA was depleted in <sup>15</sup>N relative to the microbial biomass by approximately 3.4‰. Our results show that accurate and precise isotopic measurements of C and N in DNA extracted from the soil are feasible, and that these analyses may provide powerful tools for elucidating C and N cycling processes through soil microorganisms.
2006
Barnard R, Le Roux X, Hungate BA, Cleland EE, Blankinship JC, Barthes L, Leadley PW (2006) Several components of global change alter nitrifying and denitrifying activities in an annual grassland. Functional Ecology 20(4): 557-564.
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Read Publication<span class="bullet">1. </span>The effects of global change on below-ground processes of the nitrogen (N) cycle have repercussions for plant communities, productivity and trace gas effluxes. However, the interacting effects of different components of global change on nitrification or denitrification have rarely been studied <em>in situ</em>.
<span class="bullet">2. </span>We measured responses of nitrifying enzyme activity (NEA) and denitrifying enzyme activity (DEA) to over 4 years of exposure to several components of global change and their interaction (increased atmospheric CO<sub>2</sub> concentration, temperature, precipitation and N addition) at peak biomass period in an annual grassland ecosystem. In order to provide insight into the mechanisms controlling the response of NEA and DEA to global change, we examined the relationships between these activities and soil moisture, microbial biomass C and N, and soil extractable N.
<span class="bullet">3. </span>Across all treatment combinations, NEA was decreased by elevated CO<sub>2</sub> and increased by N addition. While elevated CO<sub>2</sub> had no effect on NEA when not combined with other treatments, it suppressed the positive effect of N addition on NEA in all the treatments that included N addition. We found a significant CO<sub>2</sub>–N interaction for DEA, with a positive effect of elevated CO<sub>2</sub> on DEA only in the treatments that included N addition, suggesting that N limitation of denitrifiers may have occurred in our system. Soil water content, extractable N concentrations and their interaction explained 74% of the variation in DEA.
<span class="bullet">4. </span>Our results show that the potentially large and interacting effects of different components of global change should be considered in predicting below-ground N responses of Mediterranean grasslands to future climate changes.
Day FP, Stover DB, Pagel AL, Hungate BA, Dilustro JJ, Herbert BT, Drake BG, Hinkle CR (2006) Rapid root closure after fire limits fine root responses to elevated atmospheric CO2 in a scrub oak ecosystem in central Florida, USA. Global Change Biology 12(6): 1047-1053.
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Read PublicationElevated atmospheric carbon dioxide (CO<sub>2</sub>) often stimulates the growth of fine roots, yet there are few reports of responses of intact root systems to long-term CO<sub>2</sub> exposure. We investigated the effects of elevated CO<sub>2</sub> on fine root growth using open top chambers in a scrub oak ecosystem at Kennedy Space Center, Florida for more than 7 years. CO<sub>2</sub> enrichment began immediately after a controlled burn, which simulated the natural disturbance that occurs in this system every 10–15 years. We hypothesized that (1) root abundance would increase in both treatments as the system recovered from fire; (2) elevated CO<sub>2</sub> would stimulate root growth; and (3) elevated CO<sub>2</sub> would alter root distribution. Minirhizotron tubes were used to measure fine root length density (mm cm<sup>−2</sup>) every three months. During the first 2 years after fire recovery, fine root abundance increased in all treatments and elevated CO<sub>2</sub> significantly enhanced root abundance, causing a maximum stimulation of 181% after 20 months. The CO<sub>2</sub> stimulation was initially more pronounced in the top 10 cm and 38–49 cm below the soil surface. However, these responses completely disappeared during the third year of experimental treatment: elevated CO<sub>2</sub> had no effect on root abundance or on the depth distribution of fine roots during years 3–7. The results suggest that, within a few years following fire, fine roots in this scrub oak ecosystem reach closure, defined here as a dynamic equilibrium between production and mortality. These results further suggest that elevated CO<sub>2</sub> hastens root closure but does not affect maximum root abundance. Limitation of fine root growth by belowground resources – particularly nutrients in this nutrient-poor soil – may explain the transient response to elevated CO<sub>2</sub>.
De Graaff MA, Van Groenigen KJ, Six J, Hungate BA, van Kessel C (2006) Interactions between plant growth and soil nutrient cycling under elevated CO2: A meta‐analysis. Global Change Biology 12(11): 2077-2091.
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Read PublicationFree air carbon dioxide enrichment (FACE) and open top chamber (OTC) studies are valuable tools for evaluating the impact of elevated atmospheric CO<sub>2</sub> on nutrient cycling in terrestrial ecosystems. Using meta-analytic techniques, we summarized the results of 117 studies on plant biomass production, soil organic matter dynamics and biological N<sub>2</sub> fixation in FACE and OTC experiments. The objective of the analysis was to determine whether elevated CO<sub>2</sub> alters nutrient cycling between plants and soil and if so, what the implications are for soil carbon (C) sequestration. Elevated CO<sub>2</sub> stimulated gross N immobilization by 22%, whereas gross and net N mineralization rates remained unaffected. In addition, the soil C : N ratio and microbial N contents increased under elevated CO<sub>2</sub> by 3.8% and 5.8%, respectively. Microbial C contents and soil respiration increased by 7.1% and 17.7%, respectively. Despite the stimulation of microbial activity, soil C input still caused soil C contents to increase by 1.2% yr<sup>−1</sup>. Namely, elevated CO<sub>2</sub> stimulated overall above- and belowground plant biomass by 21.5% and 28.3%, respectively, thereby outweighing the increase in CO<sub>2</sub> respiration. In addition, when comparing experiments under both low and high N availability, soil C contents (+2.2% yr<sup>−1</sup>) and above- and belowground plant growth (+20.1% and+33.7%) only increased under elevated CO<sub>2</sub> in experiments receiving the high N treatments. Under low N availability, above- and belowground plant growth increased by only 8.8% and 14.6%, and soil C contents did not increase. Nitrogen fixation was stimulated by elevated CO<sub>2</sub> only when additional nutrients were supplied. These results suggest that the main driver of soil C sequestration is soil C input through plant growth, which is strongly controlled by nutrient availability. In unfertilized ecosystems, microbial N immobilization enhances acclimation of plant growth to elevated CO<sub>2</sub> in the long-term. Therefore, increased soil C input and soil C sequestration under elevated CO<sub>2</sub> can only be sustained in the long-term when additional nutrients are supplied.
Dijkstra P, Ishizu A, Doucett R, Hart SC, Schwartz E, Menyailo OV, Hungate BA (2006) 13C and 15N natural abundance of the soil microbial biomass. Soil Biology and Biochemistry 38(11): 3257-3266.
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Read PublicationStable isotope analysis is a powerful tool in the study of soil organic matter formation. It is often observed that more decomposed soil organic matter is <sup>13</sup>C, and especially <sup>15</sup>N-enriched relative to fresh litter and recent organic matter. We investigated whether this shift in isotope composition relates to the isotope composition of the microbial biomass, an important source for soil organic matter. We developed a new approach to determine the natural abundance C and N isotope composition of the microbial biomass across a broad range of soil types, vegetation, and climates. We found consistently that the soil microbial biomass was <sup>15</sup>N-enriched relative to the total (3.2 ‰) and extractable N pools (3.7 ‰), and <sup>13</sup>C-enriched relative to the extractable C pool (2.5 ‰). The microbial biomass was also <sup>13</sup>C-enriched relative to total C for soils that exhibited a C3-plant signature (1.6 ‰), but <sup>13</sup>C-depleted for soils with a C4 signature (−1.1 ‰). The latter was probably associated with an increase of annual C3 forbs in C4 grasslands after an extreme drought. These findings are in agreement with the proposed contribution of microbial products to the stabilized soil organic matter and may help explain the shift in isotope composition during soil organic matter formation.
Dijkstra P, Menyailo OV, Doucett RR, Hart SC, Schwartz E, Hungate BA (2006) C and N availability affects the 15N natural abundance of the soil microbial biomass across a cattle manure gradient. European Journal of Soil Science 57(4): 468-475.
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Read PublicationThe availability of C and N to the soil microbial biomass is an important determinant of the rates of soil N transformations. Here, we present evidence that changes in C and N availability affect the <sup>15</sup>N natural abundance of the microbial biomass relative to other soil N pools. We analysed the <sup>15</sup>N natural abundance signature of the chloroform-labile, extractable, NO<sub>3</sub><sup>–</sup>, NH<sub>4</sub><sup>+</sup> and soil total N pools across a cattle manure gradient associated with a water reservoir in semiarid, high-desert grassland. High levels of C and N in soil total, extractable, NO<sub>3</sub><sup>–</sup>, NH<sub>4</sub><sup>+</sup> and chloroform-labile fractions were found close to the reservoir. The δ<sup>15</sup>N value of chloroform-labile N was similar to that of extractable (organic + inorganic) N and NO<sub>3</sub><sup>–</sup> at greater C availability close to the reservoir, but was <sup>15</sup>N-enriched relative to these N-pools at lesser C availability farther away. Possible mechanisms for this variable <sup>15</sup>N-enrichment include isotope fractionation during N assimilation and dissimilation, and changes in substrate use from a less to a more <sup>15</sup>N-enriched substrate with decreasing C availability.
Goodman LF, Hungate BA (2006) Managing forests infested by spruce beetles in south-central Alaska: effects on nitrogen availability, understory biomass, and spruce regeneration. Forest ecology and management 227(3): 267-274.
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Read PublicationIn Alaska, an outbreak of spruce beetles (<em>Dendroctonus rufipennis</em>) recently infested over one million hectares of spruce (<em>Picea</em> spp.) forest. As a result, land management agencies have applied different treatments to infested forests to minimize fire hazard and economic loss and facilitate forest regeneration. In this study we investigated the effects of high-intensity burning, whole-tree harvest, whole-tree harvest with nitrogen (N) fertilization, and conventional harvest of beetle-killed stands 4 years after treatment, as well as clear-cut salvage harvest 6 years after treatment. We measured available soil ammonium and nitrate and estimated N loss from leaching using in situ cation and anion resin exchange capsules. We also assessed spruce regeneration and responses of understory plant species. Availability and losses of N did not differ among any of the management treatments. Even a substantial application of N fertilizer had no effect on N availability. Spruce regeneration significantly increased after high-intensity prescribed burning, with the number of seedlings averaging 8.9 m<sup>−2</sup> in burn plots, as compared to 0.1 m<sup>−2</sup> in plots that did not receive treatment. Biomass of the pervasive grass bluejoint (<em>Calamagrostis canadensis</em>) was significantly reduced by burning, with burn plots having 9.5% of the <em>C. canadensis</em> biomass of plots that did not receive treatment. N fertilization doubled <em>C. canadensis</em> biomass, suggesting that N fertilization without accompanying measures to control <em>C. canadensis</em> is the least viable method for promoting rapid spruce regeneration.
Hungate BA, Johnson DW, Dijkstra P, Hymus G, Stiling P, Megonigal JP, Pagel AL, Moan JL, Day F, Li J, Hinkle CR, Drake BG (2006) Nitrogen cycling during seven years of atmospheric CO2 enrichment in a scrub oak woodland. Ecology 87(1): 26-40.
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Read PublicationExperimentally increasing atmospheric CO<sub>2</sub> often stimulates plant growth and ecosystem carbon (C) uptake. Biogeochemical theory predicts that these initial responses will immobilize nitrogen (N) in plant biomass and soil organic matter, causing N availability to plants to decline, and reducing the long-term CO<sub>2</sub>-stimulation of C storage in N limited ecosystems. While many experiments have examined changes in N cycling in response to elevated CO<sub>2</sub>, empirical tests of this theoretical prediction are scarce. During seven years of postfire recovery in a scrub oak ecosystem, elevated CO<sub>2</sub> initially increased plant N accumulation and plant uptake of tracer <sup>15</sup>N, peaking after four years of CO<sub>2</sub> enrichment. Between years four and seven, these responses to CO<sub>2</sub> declined. Elevated CO<sub>2</sub> also increased N and tracer <sup>15</sup>N accumulation in the O horizon, and reduced <sup>15</sup>N recovery in underlying mineral soil. These responses are consistent with progressive N limitation: the initial CO<sub>2</sub> stimulation of plant growth immobilized N in plant biomass and in the O horizon, progressively reducing N availability to plants. Litterfall production (one measure of aboveground primary productivity) increased initially in response to elevated CO<sub>2</sub>, but the CO<sub>2</sub> stimulation declined during years five through seven, concurrent with the accumulation of N in the O horizon and the apparent restriction of plant N availability. Yet, at the level of aboveground plant biomass (estimated by allometry), progressive N limitation was less apparent, initially because of increased N acquisition from soil and later because of reduced N concentration in biomass as N availability declined. Over this seven-year period, elevated CO<sub>2</sub> caused a redistribution of N within the ecosystem, from mineral soils, to plants, to surface organic matter. In N limited ecosystems, such changes in N cycling are likely to reduce the response of plant production to elevated CO<sub>2</sub>.
Langley JA, Chapman SK, Hungate BA (2006) Ectomycorrhizal colonization slows root decomposition: the post‐mortem fungal legacy. Ecology Letters 9(8): 955-959.
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Read PublicationThe amount of carbon plants allocate to mycorrhizal symbionts exceeds that emitted by human activity annually. Senescent ectomycorrhizal roots represent a large input of carbon into soils, but their fate remains unknown. Here, we present the surprising result that, despite much higher nitrogen concentrations, roots colonized by ectomycorrhizal (EM) fungi lost only one-third as much carbon as non-mycorrhizal roots after 2 years of decomposition in a piñon pine (<em>Pinus edulis</em>) woodland. Experimentally excluding live mycorrhizal hyphae from litter, we found that live mycorrhizal hyphae may alter nitrogen dynamics, but the afterlife (litter-mediated) effects of EM fungi outweigh the influences of live fungi on root decomposition. Our findings indicate that a shift in plant allocation to mycorrhizal fungi could promote carbon accumulation in soil by this pathway. Furthermore, EM litters could directly contribute to the process of stable soil organic matter formation, a mechanism that has eluded soil scientists.
LeRoy CJ, Marks JC (2006) Litter quality, stream characteristics and litter diversity influence decomposition rates and macroinvertebrates. Freshwater Biology 51(4): 605-617.
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1. We examined the relative importance of litter quality and stream characteristics in determining decomposition rate and the macroinvertebrate assemblage living on autumn- shed leaves.
2. We compared the decomposition rates of five native riparian tree species (Populus fremontii, Alnus oblongifolia, Platanus wrightii, Fraxinus velutina and Quercus gambelii) across three south-western streams in the Verde River catchment (Arizona, U.S.A.). We also compared the decomposition of three- and five-species mixtures to that of single species to test whether plant species diversity affects rate.
3. Decomposition rate was affected by both litter quality and stream. However, litter quality accounted for most of the variation in decomposition rates. The relative importance of litter quality decreased through time, explaining 97% of the variation in the first week but only 45% by week 8. We also found that leaf mixtures decomposed more quickly than expected, when all the species included were highly labile or when the stream environment led to relatively fast decomposition.
4. In contrast to decomposition rate, differences in the invertebrate assemblage were more pronounced across streams than across leaf litter species within a stream. We also found significant differences between the invertebrate assemblage colonising leaf mixtures compared with that colonising pure species litter, indicating non-additive properties of litter diversity on stream invertebrates.
5. This study shows that leaf litter diversity has the capacity to affect in-stream decomposition rates and stream invertebrates, but that these effects depend on both litter quality and stream characteristics.
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Marks JC, Parnell R,Carter C, Dinger EC, Haden GA (2006) Interactions between geomorphology and ecosystem processes in travertine streams–implications for dam decommissioning in Fossil Creek, Arizona.. Geomorphology 779(3-4): 299-307.
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Travertine deposits of calcium carbonate can dominate channel geomorphology in streams where travertine deposition creates a distinct morphology characterized by travertine terraces, steep waterfalls, and large pools. Algae and microorganisms can facilitate travertine deposition, but how travertine affects material and energy flow in stream ecosystems is less well understood. Nearly a century of flow diversion for hydropower production has decimated the natural travertine formations in Fossil Creek, Arizona. The dam will be decommissioned in 2005. Returning carbonate-rich spring water to the natural stream channel should promote travertine deposition. How will the recovery of travertine affect the ecology of the creek? To address this question, we compared primary production, decomposition, and the abundance and diversity of invertebrates and fish in travertine and riffle/run reaches of Fossil Creek, Arizona. We found that travertine supports higher primary productivity, faster rates of leaf litter decomposition, and higher species richness of the native invertebrate assemblage. Observations from snorkeling in the stream indicate that fish density is also higher in the travertine reach. We postulate that restoring travertine to Fossil Creek will increase stream productivity, rates of litter processing, and energy flow up the food web. Higher aquatic productivity could fundamentally shift the nature of the stream from a sink to a source of energy for the surrounding terrestrial landscape.
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Menyailo OV, Hungate BA (2006) Stable isotope discrimination during soil denitrification: Production and consumption of nitrous oxide. Global Biogeochemical Cycles 20(3).
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Read Publication<span class="paraNumber">[1]</span> Measuring the stable isotope composition of nitrous oxide (N<sub>2</sub>O) evolved from soil could improve our understanding of the relative contributions of the main microbial processes (nitrification and denitrification) responsible for N<sub>2</sub>O formation in soil. However, interpretation of the isotopic data in N<sub>2</sub>O is complicated by the lack of knowledge of fractionation parameters by different microbial processes responsible for N<sub>2</sub>O production and consumption. Here we report isotopic enrichment for both nitrogen and oxygen isotopes in two stages of denitrification, N<sub>2</sub>O production and N<sub>2</sub>O reduction. We found that during both N<sub>2</sub>O production and reduction, enrichments were higher for oxygen than nitrogen. For both elements, enrichments were larger for N<sub>2</sub>O production stage than for N<sub>2</sub>O reduction. During gross N<sub>2</sub>O production, the ratio of <em>δ</em><sup>18</sup>O-to-<em>δ</em><sup>15</sup>N differed between soils, ranging from 1.6 to 2.7. By contrast, during N<sub>2</sub>O reduction, we observed a constant ratio of <em>δ</em><sup>18</sup>O-to-<em>δ</em><sup>15</sup>N with a value near 2.5. If general, this ratio could be used to estimate the proportion of N<sub>2</sub>O being reduced in the soil before escaping into the atmosphere. Because N<sub>2</sub>O-reductase enriches N<sub>2</sub>O in both isotopes, the global reduction of N<sub>2</sub>O consumption by soil may contribute to the globally observed isotopic depletion of atmospheric N<sub>2</sub>O.
Menyailo OV, Hungate BA (2006) Carbon and nitrogen stable isotopes in forest soils of Siberia. Doklady earth sciences 409(1): 747-749.
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Read Publication<div data-canvas-width="401.6631666666664">The labeled atoms of carbon and nitrogen are widely used in biology, biochemistry, and soil science. Both radioactive (14C) and stable (12C, 13C,15N) isotopes are used as such labels. Recently, an increase in mass spec-trometry sensitivity made it possible to study natural ratios of carbon (13C/12C) and nitrogen (15N/14N) stableisotopes in various biological objects. Since most bio-logical processes discriminate isotopes, i.e., use lighterisotopes for fermentative reactions [10] and leave heavier isotopes in substrate, the study of isotopic com-positions of various habitats of organisms yields infor-mation on intensity and direction of biological pro-cesses. However, the interpretation of carbon and nitro-gen isotopic compositions in complex objects, such as soil, meets difficulties because of the far too great num-ber of factors that control isotope fractionation [2, 6,11, 13]. The forest soils of the Yenisei meridian signif-icantly vary in terms of environmental factors, and thisvariability opens up possibilities to reveal the most important factors that determine carbon and nitrogen isotopic compositions of soils. In this work, the distri-bution of carbon and nitrogen isotopes in the forestsoils of the Yenisei region of Siberia was investigated for the first time. It has been shown that these isotopes are good indicators of (a) the intensity of organic matter (OM) mineralization, (b) the contribution of nitrogen fixation to the nitrogen status of ecosystems, and (c) the</div>
<div data-canvas-width="287.4446666666667">provision of ecosystems with moisture.</div>
Menyailo OV, Hungate BA (2006) Tree species and moisture effects on soil sources of N2O: Quantifying contributions from nitrification and denitrification with 18O isotopes. Journal of Geophysical Research: Biogeosciences 111(G2).
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Read Publication<span class="paraNumber">[1]</span> Nitrous oxide (N<sub>2</sub>O) is an important greenhouse gas and participates in the destruction of stratospheric ozone. Soil bacteria produce N<sub>2</sub>O through denitrification and nitrification, but these processes differ radically in substrate requirements and responses to the environment. Understanding the controls over N<sub>2</sub>O efflux from soils, and how N<sub>2</sub>O emissions may change with climate warming and altered precipitation, require quantifying the relative contributions from these groups of soil bacteria to the total N<sub>2</sub>O flux. Here we used ammonium nitrate (NH<sub>4</sub>NO<sub>3</sub>, including substrates for both processes) in which the nitrate has been enriched in the stable isotope of oxygen, <sup>18</sup>O, to partition microbial sources of N<sub>2</sub>O, arguing that a molecule of N<sub>2</sub>O carrying the <sup>18</sup>O labeled will have been produced by denitrification. We compared the influences of six common tree species on the relative contributions of nitrification and denitrification to N<sub>2</sub>O flux from soils, using soils from the Siberian afforestation experiment. We also altered soil water content, to test whether denitrification becomes a dominant source of N<sub>2</sub>O when soil water content increases. Tree species altered the proportion of nitrifier and denitrifier-derived N<sub>2</sub>O. Wetter soils produced more N<sub>2</sub>O from denitrification, though the magnitude of this effect varied among tree species. This indicates that the roles of denitrification and nitrification vary with tree species, and, that tree species influence soil responses to increased water content.
Reich PB, Hungate BA, Luo Y (2006) Carbon-nitrogen interactions in terrestrial ecosystems in response to rising atmospheric carbon dioxide. Annual Review of Ecology, Evolution, and Systematics 37: 611-636.
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Read Publication<span class="hlFld-Abstract">Interactions involving carbon (C) and nitrogen (N) likely modulate terrestrial ecosystem responses to elevated atmospheric carbon dioxide (CO<sub>2</sub>) levels at scales from the leaf to the globe and from the second to the century. In particular, response to elevated CO<sub>2</sub> may generally be smaller at low relative to high soil N supply and, in turn, elevated CO<sub>2</sub> may influence soil N processes that regulate N availability to plants. Such responses could constrain the capacity of terrestrial ecosystems to acquire and store C under rising elevated CO<sub>2</sub> levels. This review highlights the theory and empirical evidence behind these potential interactions. We address effects on photosynthesis, primary production, biogeochemistry, trophic interactions, and interactions with other resources and environmental factors, focusing as much as possible on evidence from long-term field experiments.</span>
van Groenigen KJ, Six J, Hungate BA, de Graaff M, Van Breemen N, Van Kessel C (2006) Element interactions limit soil carbon storage. Proceedings of the National Academy of Sciences 103(17): 6571-6574.
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Read PublicationRising levels of atmospheric CO<sub>2</sub> are thought to increase C sinks in terrestrial ecosystems. The potential of these sinks to mitigate CO<sub>2</sub> emissions, however, may be constrained by nutrients. By using metaanalysis, we found that elevated CO<sub>2</sub> only causes accumulation of soil C when N is added at rates well above typical atmospheric N inputs. Similarly, elevated CO<sub>2</sub> only enhances N<sub>2</sub> fixation, the major natural process providing soil N input, when other nutrients (e.g., phosphorus, molybdenum, and potassium) are added. Hence, soil C sequestration under elevated CO<sub>2</sub> is constrained both directly by N availability and indirectly by nutrients needed to support N<sub>2</sub> fixation.
2005
Barnard R, Leadley PW, Hungate BA (2005) Global change, nitrification, and denitrification: a review. Global biogeochemical cycles 19(1).
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Read Publication<span class="paraNumber">[1]</span> We reviewed responses of nitrification, denitrification, and soil N<sub>2</sub>O efflux to elevated CO<sub>2</sub>, N availability, and temperature, based on published experimental results. We used meta-analysis to estimate the magnitude of response of soil N<sub>2</sub>O emissions, nitrifying enzyme activity (NEA), denitrifying enzyme activity (DEA), and net and gross nitrification across experiments. We found no significant overall effect of elevated CO<sub>2</sub> on N<sub>2</sub>O fluxes. DEA and NEA significantly decreased at elevated CO<sub>2</sub>; however, gross nitrification was not modified by elevated CO<sub>2</sub>, and net nitrification increased. The negative overall response of DEA to elevated CO<sub>2</sub> was associated with decreased soil [NO<sub>3</sub><sup>−</sup>], suggesting that reduced availability of electron acceptors may dominate the responses of denitrification to elevated CO<sub>2</sub>. N addition significantly increased field and laboratory N<sub>2</sub>O emissions, together with gross and net nitrification, but the effect of N addition on field N<sub>2</sub>O efflux was not correlated to the amount of N added. The effects of elevated temperature on DEA, NEA, and net nitrification were not significant: The small number of studies available stress the need for more warming experiments in the field. While N addition had large effects on measurements of nitrification and denitrification, the effects of elevated CO<sub>2</sub> were less pronounced and more variable, suggesting that increased N deposition is likely to affect belowground N cycling with a magnitude of change that is much larger than that caused by elevated CO<sub>2</sub>.
Hall MC, Stiling P, Hungate BA, Drake BG, Hunter MD (2005) Effects of elevated CO2 and herbivore damage on litter quality in a scrub oak ecosystem. Journal of Chemical Ecology 31(10): 2343-2356.
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Read PublicationAtmospheric CO<sub>2</sub> concentrations have increased dramatically over the last century and continuing increases are expected to have significant, though currently unpredictable, effects on ecosystems. One important process that may be affected by elevated CO<sub>2</sub> is leaf litter decomposition. We investigated the interactions among atmospheric CO<sub>2</sub>, herbivory, and litter quality within a scrub oak community at the Kennedy Space Center, Florida. Leaf litter chemistry in 16 plots of open-top chambers was followed for 3 years; eight were exposed to ambient levels of CO<sub>2</sub>, and eight were exposed to elevated levels of CO<sub>2</sub> (ambient + 350 ppmV). We focused on three dominant oak species, <em class="EmphasisTypeItalic ">Quercus geminata</em>, <em class="EmphasisTypeItalic ">Quercus myrtifolia</em>, and <em class="EmphasisTypeItalic ">Quercus chapmanii</em>. Condensed tannin concentrations in oak leaf litter were higher under elevated CO<sub>2</sub>. Litter chemistry differed among all plant species except for condensed tannins. Phenolic concentrations were lower, whereas lignin concentrations and lignin/nitrogen ratios were higher in herbivore-damaged litter independent of CO<sub>2</sub> concentration. However, changes in litter chemistry from year to year were far larger than effects of CO<sub>2</sub> or insect damage, suggesting that these may have only minor effects on litter decomposition.
Menyailo OV, Hungate BA (2005) Tree species effects on potential production and consumption of carbon dioxide, methane, and nitrous oxide: the Siberian afforestation experiment. Tree Species Effects on Soils: Implications for Global Change 293-305.
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Read PublicationChanges in tree species composition could affect how forests produce and consume greenhouse gases, because the soil microorganisms that carry out these biogeochemical transformations are often sensitive to plant characteristics. We examined the effects of thirty years of stand development under six tree species in Siberian forests (Scots pine, spruce, arolla pine, larch, aspen and birch) on potential rates of soil CO<sub>2</sub> production, N<sub>2</sub>O-reduction and N<sub>2</sub>O production during denitrification, and CH<sub>4</sub> oxidation. Because many of these activities relate to soil N turnover, we also measured net nitrification and N mineralization. Overall, the effects of tree species were more pronounced on N<sub>2</sub>O and CH<sub>4</sub> fluxes than on CO<sub>2</sub> production. Tree species altered substrate-induced respiration (SIR) and basal respiration, but the differences were not as large as those observed for N transformations. Tree species caused similar effects on denitrification potential, net N mineralization, and net nitrification, but effects on N<sub>2</sub>O reduction were idiosyncratic, resulting in a decoupling of N<sub>2</sub>O production and reduction. CH<sub>4</sub> oxidation was affected by tree species, but these effects depended on soil moisture: increasing soil moisture enhanced CH<sub>4</sub> oxidation under some tree species but decreased it under others. If global warming causes deciduous species to replace coniferous species, our results suggest that Siberian forests would support soil microbial communities with enhanced potential to consume CH<sub>4</sub> but also to produce more N<sub>2</sub>O. Future predictions of CH<sub>4</sub> uptake and N<sub>2</sub>O efflux in boreal and temperate forests need to consider changes in tree species composition together with changes in soil moisture regimes.
2004
Flaccus K, Vleig J, Marks JC, and LeRoy CJ (2004) Restoring Fossil Creek. The Science Teacher Summer Issue, 36-40.
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At 6 A.M. on the first day of winter break, a van full of high school stu- dents and teachers set out for Fossil Creek in Arizona to conduct measurements for an ongoing environmental project (which monitors changes to the creek). The group had a long day ahead—hiking about 7 km down to Fossil Creek, wading in cold water to re- trieve samples, and carrying about 13 kg of wet sample bags back up the trail. Later that day, collected samples were analyzed in an aquatic biology lab at Northern Arizona Uni- versity (NAU). That cold, winter day ended with students saying that participating in the project was a great way to begin winter break.
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Hungate BA, Stiling PD, Dijkstra P, Johnson DW, Ketterer ME, Hymus GJ, Hinkle CR, Drake BG (2004) CO2 elicits long-term decline in nitrogen fixation. Science 304(5675): 1291-1291.
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Read PublicationRising atmospheric carbon dioxide (<em>C</em><sub>a</sub>), a product of fossil fuel burning, land-use change, and cement manufacture, is expected to cause a large carbon sink in land ecosystems, partly mitigating human-driven climate change (<a id="xref-ref-1-1" class="xref-bibr" href="http://science.sciencemag.org/content/304/5675/1291#ref-1"><em>1</em></a>). Increasing biological nitrogen fixation with rising <em>C</em><sub>a</sub> has been invoked as a means to provide the N necessary to support C accumulation (<a id="xref-ref-2-1" class="xref-bibr" href="http://science.sciencemag.org/content/304/5675/1291#ref-2"><em>2</em></a>). As in many short-term experiments (<a id="xref-ref-3-1" class="xref-bibr" href="http://science.sciencemag.org/content/304/5675/1291#ref-3"><em>3</em></a>), we found that <em>C</em><sub>a</sub> enrichment increased N fixation during the first year of treatment in an oak woodland. However, the effect declined and disappeared by the third year. <em>C</em><sub>a</sub> enrichment consistently depressed N fixation during the 5th, 6th, and 7th years of treatment. Reduced availability of the micro-nutrient molybdenum, a key constituent of nitrogenase, best explains this reduction in N fixation. Our results demonstrate how multiple element interactions can influence ecosystem responses to atmospheric change and caution against expecting increased biological N fixation to fuel terrestrial C accumulation.
Luo Y, Su BO, Currie WS, Dukes JS, Finzi A, Hartwig U, Hungate BA, McMurtrie RE, Oren RAM, Parton WJ, Pataki DE, Shaw RM, Zak DR, Field CB (2004) Progressive nitrogen limitation of ecosystem responses to rising atmospheric carbon dioxide. Bioscience 54(8): 731-739.
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Read PublicationA highly controversial issue in global biogeochemistry is the regulation of terrestrial carbon (C) sequestration by soil nitrogen (N) availability. This controversy translates into great uncertainty in predicting future global terrestrial C sequestration. We propose a new framework that centers on the concept of progressive N limitation (PNL) for studying the interactions between C and N in terrestrial ecosystems. In PNL, available soil N becomes increasingly limiting as C and N are sequestered in long-lived plant biomass and soil organic matter. Our analysis focuses on the role of PNL in regulating ecosystem responses to rising atmospheric carbon dioxide concentration, but the concept applies to any perturbation that initially causes C and N to accumulate in organic forms. This article examines conditions under which PNL may or may not constrain net primary production and C sequestration in terrestrial ecosystems. While the PNL-centered framework has the potential to explain diverse experimental results and to help researchers integrate models and data, direct tests of the PNL hypothesis remain a great challenge to the research community.
Pendall E, Bridgham S, Hanson PJ, Hungate BA, Kicklighter DW, Johnson DW, Law BE, Luo Y, Megonigal JP, Olsrud M, Ryan MG, Wan S (2004) Below‐ground process responses to elevated CO2 and temperature: a discussion of observations, measurement methods, and models. New Phytologist 162(2): 311-322.
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Read PublicationRising atmospheric CO<sub>2</sub> and temperatures are probably altering ecosystem carbon cycling, causing both positive and negative feedbacks to climate. Below-ground processes play a key role in the global carbon (C) cycle because they regulate storage of large quantities of C, and are potentially very sensitive to direct and indirect effects of elevated CO<sub>2</sub> and temperature. Soil organic matter pools, roots and associated rhizosphere organisms all have distinct responses to environmental change drivers, although availability of C substrates will regulate all the responses. Elevated CO<sub>2</sub> increases C supply below-ground, whereas warming is likely to increase respiration and decomposition rates, leading to speculation that these effects will moderate one another. However, indirect effects on soil moisture availability and nutrient supply may alter processes in unexpected directions. Detailed, mechanistic understanding and modelling of below-ground flux components, pool sizes and turnover rates is needed to adequately predict long-term, net C storage in ecosystems. In this synthesis, we discuss the current status of below-ground responses to elevated CO<sub>2</sub> and temperature and potential feedback effects, methodological challenges, and approaches to integrating models and measurements.
2003
Hungate BA, Dukes JS, Shaw MR, Luo Y, Field CB (2003) Nitrogen and climate change. Science 302(5650): 1512-1513.
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Read PublicationModels project that land ecosystems may be able take up a considerable proportion of the carbon dioxide released by human activities, thereby counteracting the anthropogenic emissions. In their Perspective, <a href="http://www.sciencemag.org/cgi/content/full/302/5650/1512"> Hungate <em>et al</em>.</a> argue that these carbon uptake estimates are too high because the models do not take other nutrients such as nitrogen into account appropriately. The authors estimate that there will not be enough nitrogen available to sustain the high carbon uptake scenarios. Nutrients other than nitrogen may also affect carbon uptake in ways not captured by most models.
Hungate BA, Naiman RJ, Apps M, Cole JJ, Moldan B, Satake K, Stewart JWB, Victoria R, Vitousek PM, Melillo JM, Field CB (2003) Disturbance and element interactions. Interactions of the major biogeochemical cycles: global change and human impacts 47-62.
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Read PublicationThis paper discusses the disturbances within and between ecosystems that result in element redistribution involving element transport in the biosphere, hydrosphere, lithosphere and atmosphere. It also discusses the disturbances (e.g. floods/landslides and vulcanism/dust storms) that redistribute and mobilize element in ratios that frequently differ from ecosystem stoichiometries, influence biogeochemical interactions on large spatial and temporal scales, and create long-lasting biological and biogeochemical legacies (e.g. widespread occurrence of N limitation of net primary production in terrestrial ecosystems). Tabulated data are given showing the effects of fire disturbance on element redistribution for a tropical forest under plantation management, for Mediterranean scrub forests, and for pine and <i>Eucalyptus</i> forests. The consequences (negative and positive feedback within ecosystem) of element redistribution are also presented. In addition, the limitations of disturbance assessment, learned views and challenges for the future are given.
Johnson DW, Hungate BA, Dijkstra P, Hymus G, Hinkle CR, Stiling P, Drake BG (2003) The effects of elevated CO2 on nutrient distribution in a fire‐adapted scrub oak forest. Ecological Applications 13(5): 1388-1399.
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Read PublicationElevated carbon dioxide (CO<sub>2</sub>) caused greater accumulation of carbon (C) and nutrients in both vegetation and O horizons over a 5-yr sampling period in a scrub oak ecosystem in Florida. Elevated CO<sub>2</sub> had no effect on any measured soil property except extractable phosphorus (P), which was lower with elevated CO<sub>2</sub> after five years. Anion and cation exchange membranes showed lower available nitrogen (N) and zinc (Zn) with elevated CO<sub>2</sub>. Soils in both elevated and ambient CO<sub>2</sub> showed decreases in total C, N, sulfur (S), and cation exchange capacity, and increases in base saturation, exchangeable Ca<sup>2+</sup>, and Mg<sup>2+</sup> over the 5-yr sampling period. We hypothesize that these soil changes were a delayed response to prescribed fire, which was applied to the site just before the initiation of the experiment. In the ambient CO<sub>2</sub> treatment, the increases in vegetation and O horizon C, N, and S were offset by the losses of soil total C, N, and S, resulting in no statistically significant net changes in ecosystem C, N, or S over time. In the elevated CO<sub>2</sub> treatment, the increases in vegetation and O horizon C content outweighed the losses in soil C, resulting in a statistically significant net increase in ecosystem C content. Nitrogen and S contents showed no statistically significant change over time in the elevated CO<sub>2</sub> treatment, however. Comparisons of vegetation contents and soil pools of potassium (K), calcium (Ca), and magnesium (Mg) suggest that a substantial proportion of these nutrients were taken up from either groundwater or deep soil horizons. This study demonstrates that changes in ecosystem C sequestration due elevated CO<sub>2</sub> or any other factor cannot be accurately assessed in the absence of data on changes in soils.
Langley JA, Dijkstra P, Drake BG, Hungate BA (2003) Ectomycorrhizal colonization, biomass, and production in a regenerating scrub oak forest in response to elevated CO2. Ecosystems 6(5): 424-430.
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Read PublicationThe effects of CO<sub>2</sub> elevation on the dynamics of fine root (FR) mass and ectomycorrhizal (EM) mass and colonization were studied in situ in a Florida scrub oak system over four years of postfire regeneration. Soil cores were taken at five dates and sorted to assess the standing crop of ectomycorrhizal and fine roots. We used ingrowth bags to estimate the effects of elevated CO<sub>2</sub> on production of EM roots and fine roots. Elevated CO<sub>2</sub> tended to increase EM colonization frequency but did not affect EM mass nor FR mass in soil cores (standing mass). However, elevated CO<sub>2</sub> strongly increased EM mass and FR mass in ingrowth bags (production), but it did not affect the EM colonization frequency therein. An increase in belowground production with unchanged biomass indicates that elevated CO<sub>2</sub> may stimulate root turnover. The CO<sub>2</sub>-stimulated increase of belowground production was initially larger than that of aboveground production. The oaks may allocate a larger portion of resources to root/mycorrhizal production in this system in elevated rather than ambient CO<sub>2</sub>.
Langley JA, Hungate BA (2003) Mycorrhizal controls on belowground litter quality. Ecology 84(9): 2302-2312.
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Read PublicationPlant productivity and ecosystem productivity are strongly influenced by nutrient availability, which is largely determined by the decomposition rate of plant litter. Belowground litter inputs (dead roots, mycorrhizae, and exudates) are larger than aboveground litterfall in many systems. Chemical quality and diameter primarily control decomposition for coarse roots, but these patterns do not hold for finer classes of roots, which are frequently colonized by mycorrhizae. Though mycorrhizal status is known to drastically alter root chemistry, morphology, life span, and exudation, it has never been explicitly considered as a factor affecting root decomposition. We hypothesize that mycorrhizal status substantially influences fine root decomposition rates.
Both ectomycorrhizal (EM) and arbuscular mycorrhizal (AM) fungi can change root properties but do so in different ways. Dominant tree species of most cold and temperate forests rely heavily on EM associations. EM fungi form massive structures that envelop fine roots. Roots infected by ectomycorrhizae have higher nitrogen concentrations than nonmycorrhizal roots, which would be expected to increase decomposition rates, but much of this nitrogen is bound in recalcitrant forms, such as chitin, so the net effect on decomposition is difficult to predict. AM fungi lack elaborate, macroscopic structures and may not alter root chemistry as profoundly.
In addition to mycorrhizal roots, external fungal hyphae can contribute significantly to ecosystem carbon budgets and thereby influence rates of soil carbon turnover. Hyphae have commonly been considered a rapidly decomposing carbon pool, though this has never been demonstrated experimentally. If hyphae are produced at the expense of rapidly decomposing root exudates, then the net effect of hyphal litter production might be to reduce soil microbial activity and overall carbon cycling rates. Based on known differences in morphology and chemistry, EM hyphae may be more recalcitrant than AM hyphae. In summary, we submit that mycorrhizal status could substantially influence fine root decomposition and soil carbon processing rates, potentially explaining some of the variation observed within and among individual plant species and ecosystems.
Li J-H, Dugas WA, Hymus GJ, Johnson DP, Hinkle CR, Drake BG, Hungate BA (2003) Direct and indirect effects of elevated CO2 on transpiration from Quercus myrtifolia in a scrub‐oak ecosystem. Global Change Biology 9(1): 96-105.
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Read PublicationElevated atmospheric carbon dioxide (C<sub>a</sub>) usually reduces stomatal conductance, but the effects on plant transpiration in the field are not well understood. Using constant-power sap flow gauges, we measured transpiration from <em>Quercus myrtifolia</em> Willd., the dominant species of the Florida scrub-oak ecosystem, which had been exposed <em>in situ</em> to elevated C<sub>a</sub> (350 µmol mol<sup><b>−</b>1</sup> above ambient) in open-top chambers since May 1996. Elevated C<sub>a</sub> reduced average transpiration per unit leaf area by 37%, 48% and 49% in March, May and October 2000, respectively. Temporarily reversing the C<sub>a</sub> treatments showed that at least part of the reduction in transpiration was an immediate, reversible response to elevated C<sub>a</sub>. However, there was also an apparent indirect effect of C<sub>a</sub> on transpiration: when transpiration in all plants was measured under common C<sub>a</sub>, transpiration in elevated C<sub>a</sub>-grown plants was lower than that in plants grown in normal ambient C<sub>a</sub>. Results from measurements of stomatal conductance (g<sub>s</sub>), leaf area index (LAI), canopy light interception and correlation between light and g<sub>s</sub> indicated that the direct, reversible C<sub>a</sub> effect on transpiration was due to changes in g<sub>s</sub> caused by C<sub>a</sub>, and the indirect effect was caused mainly by greater self-shading resulting from enhanced LAI, not from stomatal acclimation. By reducing light penetration through the canopy, the enhanced self-shading at elevated C<sub>a</sub> decreased stomatal conductance and transpiration of leaves at the middle and bottom of canopy. This self-shading mechanism is likely to be important in ecosystems where LAI increases in response to elevated C<sub>a</sub>.
Menyailo OV, Hungate BA (2003) Interactive effects of tree species and soil moisture on methane consumption. Soil Biology and Biochemistry 35(4): 625-628.
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Read PublicationMethane consumption by temperate forest soils is a major sink for this important greenhouse gas, but little is known about how tree species influence CH<sub>4</sub> uptake by soils. Here, we show that six common tree species in Siberian boreal and temperate forests significantly affect potential CH<sub>4</sub> consumption in laboratory microcosms. Overall, soils under hardwood species (aspen and birch) consumed CH<sub>4</sub> at higher rates than soils under coniferous species and grassland. While NH<sub>4</sub><sup>+</sup> addition often reduces CH<sub>4</sub> uptake, we found no effect of NH<sub>4</sub><sup>+</sup> addition, possibly because of the relatively high ratio of CH<sub>4</sub>-to-NH<sub>4</sub><sup>+</sup> in our incubations. The effects of soil moisture strongly depended on plant species. An increase in soil moisture enhanced CH<sub>4</sub> consumption in soils under spruce but had the opposite effect under Scots pine and larch. Under other species, soil moisture did not affect CH<sub>4</sub> consumption. These results could be explained by specific responses of different groups of CH<sub>4</sub>-oxidizing bacteria to elevated moisture.
Menyailo OV, Hungate BA, Lehmann J, Gebauer G, Zech W (2003) Tree species of the central Amazon and soil moisture alter stable isotope composition of nitrogen and oxygen in nitrous oxide evolved from soil. Isotopes in environmental and health studies 39(1): 41-52.
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Read PublicationThe use of stable isotopes of N and O in N<sub>2</sub>O has been proposed as a way to better constrain the global budget of atmospheric N<sub>2</sub>O and to better understand the relative contributions of the main microbial processes (nitrification and denitrification) responsible for N<sub>2</sub>O formation in soil. This study compared the isotopic composition of N<sub>2</sub>O emitted from soils under different tree species in the Brazilian Amazon. We also compared the effect of tree species with that of soil moisture, as we expected the latter to be the main factor regulating the proportion of nitrifier- and denitrifier-derived N<sub>2</sub>O and, consequently, isotopic signatures of N<sub>2</sub>O. Tree species significantly affected <i>δ</i> <sup>15</sup>N in nitrous oxide. However, there was no evidence that the observed variation in <i>δ</i> <sup>15</sup>N in N<sub>2</sub>O was determined by varying proportions of nitrifier- vs. denitrifier-derived N<sub>2</sub>O. We submit that the large variation in <i>δ</i> <sup>15</sup>N-N<sub>2</sub>O is the result of competition between denitrifying and immobilizing microorganisms for NO 3 m . In addition to altering <i>δ</i> <sup>15</sup>N-N<sub>2</sub>O, tree species affected net rates of N<sub>2</sub>O emission from soil in laboratory incubations. These results suggest that tree species contribute to the large isotopic variation in N<sub>2</sub>O observed in a range tropical forest soils. We found that soil water affects both <sup>15</sup>N and <sup>18</sup>O in N<sub>2</sub>O, with wetter soils leading to more depleted N<sub>2</sub>O in both <sup>15</sup>N and <sup>18</sup>O. This is likely caused by a shift in biological processes for <sup>15</sup>N and possible direct exchange of <sup>18</sup>O between H<sub>2</sub>O and N<sub>2</sub>O.
2002
Dijkstra P, Hymus G, Colavito D, Vieglais DA, Cundari CM, Johnson DP, Hungate BA, Hinkle C, Drake BG (2002) Elevated atmospheric CO2 stimulates aboveground biomass in a fire‐regenerated scrub‐oak ecosystem. Global Change Biology 8(1): 90-103.
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Read PublicationThe effect of elevated atmospheric CO<sub>2</sub> concentration (<em>C</em><sub>a</sub>) on the aboveground biomass of three oak species, <em>Quercus myrtifolia</em>, <em>Q. geminata</em>, and <em>Q. chapmanii</em>, was estimated nondestructively using allometric relationships between stem diameter and aboveground biomass after four years of experimental treatment in a naturally fire-regenerated scrub-oak ecosystem. After burning a stand of scrub-oak vegetation, re-growing plants were exposed to either current ambient (379 µL L<sup>−1</sup> CO<sub>2</sub>) or elevated (704 µL L<sup>−1</sup> CO<sub>2</sub>) <em>C</em><sub>a</sub> in 16 open-top chambers over a four-year period, and measurements of stem diameter were carried out annually on all oak shoots within each chamber. Elevated <em>C</em><sub>a</sub> significantly increased aboveground biomass, expressed either per unit ground area or per shoot; elevated <em>C</em><sub>a</sub> had no effect on shoot density. The relative effect of elevated <em>C</em><sub>a</sub> on aboveground biomass increased each year of the study from 44% (May 96–Jan 97), to 55% (Jan 97–Jan 98), 66% (Jan 98–Jan 99), and 75% (Jan 99–Jan 00). The effect of elevated <em>C</em><sub>a</sub> was species specific: elevated <em>C</em><sub>a</sub> significantly increased aboveground biomass of the dominant species, <em>Q. myrtifolia</em>, and tended to increase aboveground biomass of <em>Q. chapmanii</em>, but had no effect on aboveground biomass of the subdominant, <em>Q. geminata</em>. These results show that rising atmospheric CO<sub>2</sub> has the potential to stimulate aboveground biomass production in ecosystems dominated by woody species, and that species-specific growth responses could, in the long term, alter the composition of the scrub-oak community.
Dukes JS, Hungate BA (2002) Elevated carbon dioxide and litter decomposition in California annual grasslands: which mechanisms matter?. Ecosystems 5(2): 171-183.
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Read PublicationTo date, most research that has examined the effect of elevated atmospheric carbon dioxide concentration ([CO<sub>2</sub>]) on litter decomposition has focused on changes in the leaf litter quality of individual species. Results from California grasslands indicate that other CO<sub>2</sub> responses may have greater consequences for decomposition rates. For instance, CO<sub>2</sub>-driven changes in either species dominance or patterns of biomass allocation would alter both the quality and the position of grassland litter. We review the results from studies in California grasslands to identify the mechanisms that affect grassland litter decomposition. We use a simple calculation that integrates the results of two studies to identify three mechanisms that have the potential to substantially alter decomposition rates as the atmospheric [CO<sub>2</sub>] rises.
Hungate BA, Reichstein M, Dijkstra P, Johnson D, Hymus G, Tenhunen JD, Hinkle CR, Drake BG (2002) Evapotranspiration and soil water content in a scrub‐oak woodland under carbon dioxide enrichment. Global Change Biology 8(3): 289-298.
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Read PublicationLeaf conductance often decreases in response to elevated atmospheric CO<sub>2</sub> concentration (<em>C</em><sub>a</sub>) potentially leading to changes in hydrology. We describe the hydrological responses of Florida scrub oak to elevated <em>C</em><sub>a</sub> during an eight-month period two years after <em>C</em><sub>a</sub> manipulation began. Whole-chamber gas exchange measurements revealed a consistent reduction in evapotranspiration in response to elevated <em>C</em><sub>a</sub>, despite an increase in leaf area index (LAI). Elevated <em>C</em><sub>a</sub> also increased surface soil water content, but xylem water deuterium measurements show that the dominant oaks in this system take up most of their water from the water table (which occurs at a depth of 1.5–3 m), suggesting that the water savings in elevated <em>C</em><sub>a</sub> in this system are primarily manifested as reduced water uptake at depth. Extrapolating these results to larger areas requires considering a number of processes that operate on scales beyond these accessible in this field experiment. Nevertheless, these results demonstrate the potential for reduced evapotranspiration and associated changes in hydrology in ecosystems dominated by woody vegetation in response to elevated <em>C</em><sub>a</sub>.
Hymus GJ, Snead TG, Johnson DP, Hungate BA, Drake BG (2002) Acclimation of photosynthesis and respiration to elevated atmospheric CO2 in two Scrub Oaks. Global Change Biology 8(4): 317-328.
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Read PublicationFor two species of oak, we determined whether increasing atmospheric CO<sub>2</sub> concentration (<em>C</em><sub>a</sub>) would decrease leaf mitochondrial respiration (R) directly, or indirectly owing to their growth in elevated <em>C</em><sub>a</sub>, or both. In particular, we tested whether acclimatory decreases in leaf-Rubisco content in elevated <em>C</em><sub>a</sub> would decrease R associated with its maintenance. This hypothesis was tested in summer 2000 on sun and shade leaves of <em>Quercus myrtifolia</em> Willd. and <em>Quercus geminata</em> Small. We also measured R on five occasions between summer 1999 and 2000 on leaves of <em>Q. myrtifolia</em>. The oaks were grown in the field for 4 years, in either current ambient or elevated (current ambient + 350 µmol mol<sup>−1</sup>) <em>C</em><sub>a</sub>, in open-top chambers (OTCs). For <em>Q. myrtifolia</em>, an increase in <em>C</em><sub>a</sub> from 360 to 710 µmol mol<sup>−1</sup> had no direct effect on R at any time during the year. In April 1999, R in young <em>Q. myrtifolia</em> leaves was significantly higher in elevated <em>C</em><sub>a</sub>—the only evidence for an indirect effect of growth in elevated <em>C</em><sub>a</sub>. Leaf R was significantly correlated with leaf nitrogen (N) concentration for the sun and shade leaves of both the species of oak. Acclimation of photosynthesis in elevated <em>C</em><sub>a</sub> significantly reduced maximum RuBP-saturated carboxylation capacity (<em>V</em><sub>c max</sub>) for both the sun and shade leaves of only <em>Q. geminata</em>. However, we estimated that only 11–12% of total leaf N was invested in Rubisco; consequently, acclimation in this plant resulted in a small effect on N and an insignificant effect on R. In this study measurements of respiration and photosynthesis were made on material removed from the field; this procedure had no effect on gas exchange properties. The findings of this study were applicable to R expressed either per unit leaf area or unit dry weight, and did not support the hypothesis that elevated <em>C</em><sub>a</sub> decreases R directly, or indirectly owing to acclimatory decreases in Rubisco content.
Langley J, Drake B, Hungate B (2002) Extensive belowground carbon storage supports roots and mycorrhizae in regenerating scrub oaks. Oecologia 131(4): 542-548.
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Read PublicationPortions of a regenerating scrub oak ecosystem were enclosed in open-top chambers and exposed to elevated CO<sub>2</sub>. The distinct <sup>13</sup>C signal of the supplemental CO<sub>2</sub> was used to trace the rate of C integration into various ecosystem components. Oak foliage, stems, roots and ectomycorrhizae were sampled over 3 years and were analyzed for <sup>13</sup>C composition. The aboveground tissue <sup>13</sup>C equilibrated to the novel <sup>13</sup>C signal in the first season, while the belowground components displayed extremely slow integration of the new C. Roots taken from ingrowth cores showed that 33% of the C in newly formed roots originated from a source other than recent photosynthesis inside the chamber. In this highly fire-prone system, the oaks re-establish primarily by resprouting from large rhizomes. Remobilization from belowground C stores may support fine roots and mycorrhizae for several years into stand re-establishment and, therefore, may explain why belowground tissues contain less of the new photosynthate than expected. Though it has been shown that long-term cycles of C storage are theoretically advantageous for plants in systems with frequent and severe disturbances, such patterns have not been previously examined in wild systems.
Menyailo OV, Hungate BA, Zech W (2002) The effect of single tree species on soil microbial activities related to C and N cycling in the Siberian artificial afforestation experiment. Plant and Soil 242(2): 183-196.
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Read PublicationThe effects of grassland conversion to forest vegetation and of individual tree species on microbial activity in Siberia are largely unstudied. Here, we examined the effects of the six most commonly dominant tree species in Siberian forests (Scots pine, spruce, Arolla pine, larch, aspen and birch) on soil C and N mineralization, N<sub>2</sub>O-reduction and N<sub>2</sub>O production during denitrification 30 years after planting. We also documented the effect of grassland conversion to different tree species on microbial activities at different soil depths and their relationships to soil chemical properties. The effects of tree species and grassland conversion were more pronounced on N than on C transformations. Tree species and grassland conversion did significantly alter substrate-induced respiration (SIR) and basal respiration, but the differences were not as large as those observed for N transformations. Variances in SIR and basal respiration within species were markedly lower than those in N transformations. Net N mineralization, net nitrification, and denitrification potential were highest under Arolla pine and larch, intermediate under deciduous aspen and birch, and lowest beneath spruce and Scots pine. Tree species caused similar effects on denitrification potential, net N mineralization, and net nitrification, but effects on N<sub>2</sub>O reduction rate were idiosyncratic, indicating a decoupling of N<sub>2</sub>O production and reduction. We predict that deciduous species should produce more N<sub>2</sub>O in the field than conifers, and that Siberian forests will produce more N<sub>2</sub>O if global climate change alters tree species composition. Basal respiration and SIR showed inverse responses to tree species: when basal respiration increased in response to a given tree species, SIR declined. SIR may have been controlled by NH<sub>4</sub> <sup>+</sup> availability and related therefore to N mineralization, which was negatively affected by grassland conversion. Basal respiration appeared to be less limited by NH<sub>4</sub> <sup>+</sup> and controlled mostly by readily available organic C (DOC), which was higher in concentration under forests than in grassland and therefore basal respiration was higher in forested soils. We conclude that in the Siberian artificial afforestation experiment, soil C mineralization was not limited by N.
Stiling P, Cattell M, Moon DC, Rossi A, Hungate BA, Hymus G, Drake B (2002) Elevated atmospheric CO2 lowers herbivore abundance, but increases leaf abscission rates. Global Change Biology 8(7): 658-667.
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Read PublicationIncreased levels of atmospheric carbon dioxide (CO<sub>2</sub>) are likely to affect the trophic relationships that exist between plants, their herbivores and the herbivores' natural enemies. This study takes advantage of an open-top CO<sub>2</sub> fertilization experiment in a Florida scrub oak community at Kennedy Space Center, Florida, consisting of eight chambers supplied with ambient CO<sub>2</sub> (360 ppm) and eight chambers supplied with elevated CO<sub>2</sub> (710 ppm). We examined the effects of elevated CO<sub>2</sub> on herbivore densities and levels of leaf consumption, rates of herbivore attack by natural enemies and effects on leaf abscission. Cumulative levels of herbivores and herbivore damage were significantly lower in elevated CO<sub>2</sub> than in ambient CO<sub>2</sub>. This may be because leaf nitrogen levels are lower in elevated CO<sub>2</sub>. More herbivores die of host plant-induced death in elevated CO<sub>2</sub> than in ambient CO<sub>2</sub>. Attack rates of herbivores by parasitoids are also higher in elevated CO<sub>2</sub>, possibly because herbivores need to feed for a longer time in order to accrue sufficient nitrogen (N), thus exposing themselves longer to natural enemies. Insect herbivores cause an increase in abscission rates of leaves throughout the year. Because of the lower insect density in elevated CO<sub>2</sub>, we thought, abscission rates would be lower in these chambers. However, abscission rates were significantly higher in elevated CO<sub>2</sub>. Thus, the direct effects of elevated CO<sub>2</sub> on abscission are greater than the indirect effects on abscission mediated via lower insect densities. A consequence of increased leaf abscission in elevated CO<sub>2</sub> is that nutrient deposition rates to the soil surface are accelerated.
2001
Cardon ZG, Hungate BA, Cambardella CA, Chapin FS, Field CB, Holland EA, Mooney HA (2001) Contrasting effects of elevated CO 2 on old and new soil carbon pools. Soil Biology and Biochemistry 33(3): 365-373.
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Read PublicationSoil organic carbon (SOC) is the largest reservoir of organic carbon in the terrestrial biosphere. Though the influence of increasing atmospheric CO<sub>2</sub> on net primary productivity, on the flow of newly fixed carbon belowground, and on the quality of new plant litter in ecosystems has been examined, indirect effects of increased CO<sub>2</sub> on breakdown of large SOC pools already in ecosystems are not well understood. We found that exposure of California grassland communities to elevated CO<sub>2</sub> retarded decomposition of older SOC when mineral nutrients were abundant, thus increasing the turnover time of SOC already in the system. Under elevated CO<sub>2</sub>, soil microorganisms appeared to shift from consuming older SOC to utilizing easily degraded rhizodeposits derived from increased root biomass. In contrast to this increased retention of stabilized older SOC under elevated CO<sub>2</sub>, movement of newly fixed carbon from roots to stabilized SOC pools was retarded; though root biomass increased under elevated CO<sub>2</sub>, new carbon in mineral-bound pools decreased. These contrasting effects of elevated CO<sub>2</sub> on dynamics of old and new soil carbon pools contribute to a new soil carbon equilibrium that could profoundly affect long-term net carbon movement between terrestrial ecosystems and the atmosphere.
Hungate BA, Marks JC (2001) Terrestrial and freshwater ecosystems: impacts of global change. Encyclopaedia of Global Environmental Change 2: 122-134.
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Download .PDFHumans are altering the environment in many ways. While local environmental damage (landfills, oil spills, urban smog) is still prevalent, we now also realize that human activities are altering the Earth System as a whole, that our environmental crisis has become truly global.
Johnson DW, Hungate B, Dijkstra P, Hymus G, Drake B (2001) Effects of elevated carbon dioxide on soils in a Florida scrub oak ecosystem. Journal of Environmental Quality 30(2): 501-507.
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Read PublicationThe results of a 3-yr study on the effects of elevated CO<sub>2</sub> on soil N and P, soil pCO<sub>2</sub>, and calculated CO<sub>2</sub> efflux in a fire-regenerated Florida scrub oak ecosystem are summarized. We hypothesized that elevated CO<sub>2</sub> would cause (i) increases in soil pCO<sub>2</sub> and soil respiration and (ii) reduced levels of soil-available N and P. The effects of elevated CO<sub>2</sub> on soil N availability differed according to the method used. Results of resin lysimeter collections and anion exchange membrane tests in the field showed reduced NO<sup>−</sup> <sub>3</sub> in soils in Years 1 and 3. On the other hand, re-analysis of homogenized, buried soil bags after 1 yr suggested a relative increase in N availability (lower C to N ratio) under elevated CO<sub>2</sub> In the case of P, the buried bags and membranes suggested a negative effect of CO<sub>2</sub> on P during the first year; this faded over time, however, as P availability declined overall, probably in response to P uptake. Elevated CO<sub>2</sub> had no effect on soil pCO<sub>2</sub> or calculated soil respiration at any time, further suggesting that plant rather than microbial uptake was the primary factor responsible for the observed changes in N and P availability with elevated CO<sub>2</sub>
2000
Hungate BA, Jaeger III CH, Gamara G, Chapin III FS, Field CB (2000) Soil microbiota in two annual grasslands: responses to elevated atmospheric CO2. Oecologia 124(4): 589-598.
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Read PublicationWe measured soil bacteria, fungi, protozoa, nematodes, and biological activity in serpentine and sandstone annual grasslands after 4 years of exposure to elevated atmospheric CO<sub>2</sub>. Measurements were made during the early part of the season, when plants were in vegetative growth, and later in the season, when plants were approaching their maximum biomass. In general, under ambient CO<sub>2</sub>, bacterial biomass, total protozoan numbers, and numbers of bactivorous nematodes were similar in the two grasslands. Active and total fungal biomasses were higher on the more productive sandstone grassland compared to the serpentine. However, serpentine soils contained nearly twice the number of fungivorous nematodes compared to the sandstone, perhaps explaining the lower standing crop of fungal biomass in the serpentine and suggesting higher rates of energy flow through the fungal-based soil food web. Furthermore, root biomass in the surface soils of these grasslands is comparable, but the serpentine contains 6 times more phytophagous nematodes compared to the sandstone, indicating greater below-ground grazing pressure on plants in stressful serpentine soils. Elevated CO<sub>2</sub> increased the biomass of active fungi and the numbers of flagellates in both grasslands during the early part of the season and increased the number of phytophagous nematodes in the serpentine. Elevated CO<sub>2</sub> had no effect on the total numbers of bactivorous or fungivorous nematodes, but decreased the diversity of the nematode assemblage in the serpentine at both sampling dates. Excepting this reduction in nematode diversity, the effects of elevated CO<sub>2</sub> disappeared later in the season as plants approached their maximum biomass. Elevated CO<sub>2</sub> had no effect on total and active bacterial biomass, total fungal biomass, or the total numbers of amoebae and ciliates in either grassland during either sampling period. However, soil metabolic activity was higher in the sandstone grassland in the early season under elevated CO<sub>2</sub>, and elevated CO<sub>2</sub> altered the patterns of use of individual carbon substrates in both grasslands at this time. Rates of substrate use were also significantly higher in the sandstone, indicating increased bacterial metabolic activity. These changes in soil microbiota are likely due to an increase in the flux of carbon from roots to soil in elevated CO<sub>2</sub>, as has been previously reported for these grasslands. Results presented here suggest that some of the carbon distributed below ground in response to elevated CO<sub>2</sub> affects the soil microbial food web, but that these effects may be more pronounced during the early part of the growing season.
1999
Hungate BA (1999) Ecosystem responses to rising atmospheric CO2: feedbacks through the nitrogen cycle. Carbon dioxide and environmental stress 265-285.
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Read PublicationThis chapter discusses the mechanisms through which elevated CO2 can cause changes in soil nitrogen cycling. Rising atmospheric CO2 could alter soil nitrogen (N) cycling, shaping the responses of many terrestrial ecosystems to elevated CO2. Increased carbon input to soil through increased root growth, altered litter quality, and increased soil water content through decreased plant water use in elevated CO2 can all affect soil N transformations and thus, N availability to plants. Nitrogen limits net primary productivity (NPP) in many terrestrial ecosystems, so changes in N availability to plants will influence NPP in an elevated CO2 environment. Furthermore, changes in NPP will alter carbon uptake by the terrestrial biosphere, and thus, feed back to rising atmospheric CO2. Elevated CO2 could also influence the processes that regulate N inputs to and losses from ecosystems —N fixation, gaseous N losses, and N leaching. Such changes could alter ecosystem nitrogen stocks and thus, nitrogen available to support NPP. Additionally, soil emissions of N20 contribute to the greenhouse effect and stratospheric ozone destruction, and emissions of NOx contribute to photochemical smog and acid rain. Thus, by altering soil nitrogen cycling, elevated CO2 could cause other changes in atmospheric chemistry.
Hungate BA, Dijkstra P, Johnson D, Hinkle CR, Drake B (1999) Elevated CO2 increases nitrogen fixation and decreases soil nitrogen mineralization in Florida scrub oak. Global Change Biology 5(7): 781-789.
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Read PublicationWe report changes in nitrogen cycling in Florida scrub oak in response to elevated atmospheric CO<sub>2</sub> during the first 14 months of experimental treatment. Elevated CO<sub>2</sub> stimulated above-ground growth, nitrogen mass, and root nodule production of the nitrogen-fixing vine, <em>Galactia elliottii</em> Nuttall. During this period, elevated CO<sub>2</sub> reduced rates of gross nitrogen mineralization in soil, and resulted in lower recovery of nitrate on resin lysimeters. Elevated CO<sub>2</sub> did not alter nitrogen in the soil microbial biomass, but increased the specific rate of ammonium immobilization (NH<sub>4</sub><sup>+</sup> immobilized per unit microbial N) measured over a 24-h period. Increased carbon input to soil through greater root growth combined with a decrease in the quality of that carbon in elevated CO<sub>2</sub> best explains these changes.
These results demonstrate that atmospheric CO<sub>2</sub> concentration influences both the internal cycling of nitrogen (mineralization, immobilization, and nitrification) as well as the processes that regulate total ecosystem nitrogen mass (nitrogen fixation and nitrate leaching) in Florida coastal scrub oak. If these changes in nitrogen cycling are sustained, they could cause long-term feedbacks to the growth responses of plants to elevated CO<sub>2</sub>. Greater nitrogen fixation and reduced leaching could stimulate nitrogen-limited plant growth by increasing the mass of labile nitrogen in the ecosystem. By contrast, reduced nitrogen mineralization and increased immobilization will restrict the supply rate of plant-available nitrogen, potentially reducing plant growth. Thus, the net feedback to plant growth will depend on the balance of these effects through time.
Stiling P, Rossi AM, Hungate B, Dijkstra P, Hinkle CR, Knott WM, Drake B (1999) Decreased leaf‐miner abundance in elevated CO2: Reduced leaf quality and increased parasitoid attack. Ecological Applications 9(1): 240-244.
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Read PublicationMost studies on the effects of elevated CO<sub>2</sub> have focused on the effects on plant growth and ecosystem processes. Fewer studies have examined the effects of elevated CO<sub>2</sub> on herbivory, and of these, most have examined feeding rates in laboratory conditions. Our study takes advantage of an open-top CO<sub>2</sub> fertilization study in a Florida scrub-oak community to examine the effects of elevated CO<sub>2</sub> on herbivore densities, herbivore feeding rates, and levels of attack of herbivores by natural enemies.
Higher atmospheric CO<sub>2</sub> concentration reduced plant foliar nitrogen concentrations, decreased abundance of leaf-mining insect herbivores, increased per capita leaf consumption by leafminers, and increased leafminer mortality. As suggested by other authors, reduced foliar quality contributed to the increase in herbivore mortality, but only partly. The major factor increasing mortality was higher attack rate by parasitoids. Thus increasing CO<sub>2</sub> concentrations may reduce the survivorship of insect herbivores directly, by reducing plant quality, but also indirectly, by changing herbivore feeding and eliciting greater top-down pressure from natural enemies.
1997
Franck VM, Hungate BA, Chapin III FS, Field CB (1997) Decomposition of litter produced under elevated CO2: dependence on plant species and nutrient supply. Biogeochemistry 36(3): 223-237.
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Read PublicationWe investigated the effect of CO<sub>2</sub> concentration and soilnutrient availability during growth on the subsequent decomposition andnitrogen (N) release from litter of four annual grasses that differ inresource requirements and native habitat. Vulpia microstachys isa native grass found on California serpentine soils, whereas Avenafatua, Bromus hordaceus, and Lolium multiflorum areintroduced grasses restricted to more fertile sandstone soils (Hobbs & Mooney 1991). Growth in elevated CO<sub>2</sub> altered litter C:N ratio,decomposition, and N release, but the direction and magnitude of thechanges differed among plant species and nutrient treatments. ElevatedCO<sub>2</sub> had relatively modest effects on C:N ratio of litter,increasing this ratio in Lolium roots (and shoots at high nutrients),but decreasing C:N ratio in Avena shoots. Growth of plants underelevated CO<sub>2</sub> decreased the decomposition rate of Vulpialitter, but increased decomposition of Avena litter from the high-nutrient treatment. The impact of elevated CO<sub>2</sub> on N loss fromlitter also differed among species, with Vulpia litter from high-CO<sub>2</sub> plants releasing N more slowly than ambient-CO<sub>2</sub>litter, whereas growth under elevated CO<sub>2</sub> caused increased Nloss from Avena litter. CO<sub>2</sub> effects on N release in Lolium and Bromus depended on the nutrient regime in whichplants were grown. There was no overall relationship between litter C:Nratio and decomposition rate or N release across species and treatments.Based on our study and the literature, we conclude that the effects ofelevated CO<sub>2</sub> on decomposition and N release from litter arehighly species-specific. These results do not support the hypothesis thatCO<sub>2</sub> effects on litter quality consistently lead to decreasednutrient availability in nutrient-limited ecosystems exposed to elevatedCO<sub>2</sub>.
Garland JL, Cook KL, Loader CA, Hungate BA (1997) The influence of microbial community structure and function on community-level physiological profiles. Microbial Communities 171-183.
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Read PublicationPatterns of carbon source utilization, or community-level physiological profiles (CLPP), produced from direct incubation of environmental samples in BIOLOG microplates can consistently discriminate spatial and temporal gradients within microbial communities. While the resolving power of the assay appears significant, the basis for the differences in the patterns of sole carbon source utilization among communities remains unclear. Carbon source utilization as measured in this assay is a measure of functional potential, rather than <em class="EmphasisTypeItalic ">in situ</em> activity, since enrichment occurs over the course of incubation, which can range from 24 to 72 hours (or even longer) depending on inoculum density. The functional profile of a community could be an indicator of carbon source availability and concomitant selection for specific functional types of organisms. A more limited view of the profile is as a composite descriptor of the microbial community composition without any ecologically relevant functional information. We manipulated microbial community structure and function in laboratory microcosms to evaluate their influence on CLPP. The structure of rhizosphere communities was controlled by inoculating axenic plants (wheat and potato) with different mixed species (non-gnotobiotic) inocula. Inoculum source influenced CLPP more strongly than plant type, indicating that CLPP primarily reflected differences in microbial community structure than function. In order to more specifically examine the influence of microbial function on CLPP, specific carbon sources in the BIOLOG plates (asparagine and acetate) were added to a continuously stirred tank reactor (CSTR) containing a mixed community of microorganisms degrading plant material. Daily additions of these carbon sources at levels up to 50% of the total respired carbon in the bioreactor caused significant changes in overall CLPP, but caused no, or minor, increases in the specific response of these substrates in the plates. These studies indicate that the functional relevance of CLPP should be interpreted with caution.
Hungate BA, Holland EA, Jackson RB, Chapin FS, Mooney HA, Field CB (1997) The fate of carbon in grasslands under carbon dioxide enrichment. Nature 388(6642): 576-579.
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Read PublicationThe concentration of carbon dioxide (CO<sub>2</sub>) in the Earth's atmosphere is rising rapidly<sup><a href="http://www.nature.com/nature/journal/v388/n6642/full/388576a0.html#B1">1</a></sup>, with the potential to alter many ecosystem processes. Elevated CO<sub>2</sub> often stimulates photosynthesis<sup><a href="http://www.nature.com/nature/journal/v388/n6642/full/388576a0.html#B2">2</a></sup>, creating the possibility that the terrestrial biosphere will sequester carbon in response to rising atmospheric CO<sub>2</sub> concentration, partly offsetting emissions from fossil-fuel combustion, cement manufacture, and deforestation<sup><a href="http://www.nature.com/nature/journal/v388/n6642/full/388576a0.html#B3">3</a></sup>,<sup><a href="http://www.nature.com/nature/journal/v388/n6642/full/388576a0.html#B4">4</a></sup>. However, the responses of intact ecosystems to elevated CO<sub>2</sub> concentration, particularly the below-ground responses, are not well understood. Here we present an annual budget focusing on below-ground carbon cycling for two grassland ecosystems exposed to elevated CO<sub>2</sub> concentrations. Three years of experimental CO<sub>2</sub> doubling increased ecosystem carbon uptake, but greatly increased carbon partitioning to rapidly cycling carbon pools below ground. This provides an explanation for the imbalance observed in numerous CO<sub>2</sub> experiments, where the carbon increment from increased photosynthesis is greater than the increments in ecosystem carbon stocks. The shift in ecosystem carbon partitioning suggests that elevated CO<sub>2</sub> concentration causes a greater increase in carbon cycling than in carbon storage in grasslands.
Hungate BA, Jordan TE, Jackson RB, Drake BG (1997) Atmospheric nitrogen deposition. Science 275(5301): 737-741.
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Read Publication<p id="p-1">David A. Wedin and David Tilman (Reports, 6 Dec.,<a href="http://science.sciencemag.org/lookup/volpage/275/1720">p 1720</a>) show that increased nitrogen inputs to terrestrial ecosystems might cause smaller increases in the capacity of those ecosystems to store carbon than expected. Their findings are important because nitrogen inputs have increased dramatically over the past decades through fertilizer production, cultivation of nitrogen-fixing legumes, and production of oxides of nitrogen associated with fossil-fuel burning (<a id="xref-ref-1-1" class="xref-bibr" href="http://science.sciencemag.org/content/275/5301/737.5#ref-1">1</a>). However, the simultaneous increase in atmospheric carbon dioxide (CO<sub>2</sub>) concentrations caused by burning fossil fuels is likely to at least partially counteract the processes that limited carbon storage in Wedin and Tilman's experiment. CO<sub>2</sub> enrichment generally increases the amount of carbon fixed by plants per unit of nitrogen taken up from the soil, particularly in carbon-3 (C<sup>3</sup>) species (<a id="xref-ref-2-1" class="xref-bibr" href="http://science.sciencemag.org/content/275/5301/737.5#ref-2">2</a>) such as those that invaded their nitrogen-enriched plots. Compared with the C<sup>4</sup> species that thrived before nitrogen was added, the invading C<sup>3</sup> species have relatively lower C-to-N ratios, limiting the amount of carbon stored in response to nitrogen input. However, with elevated CO<sub>2</sub> tending to increase the C-to-N ratio of these C<sup>3</sup> plants, N and CO<sub>2</sub> enrichment in concert would likely cause greater C storage than observed by Wedin and Tilman.</p>
<p id="p-2">Rising atmospheric CO<sub>2</sub> may also increase N inputs to terrestrial ecosystems, amplifying the direct human impact on the N cycle. CO<sub>2</sub> enrichment often increases the growth of plants housing N-fixing bacteria in their roots, and this stimulation is relatively larger than non-N-fixing plants (<a id="xref-ref-3-1" class="xref-bibr" href="http://science.sciencemag.org/content/275/5301/737.5#ref-3">3</a>). Thus, in addition to the direct anthropogenic stimulation of N inputs to terrestrial ecosystems through agriculture and fossil-fuel burning (<a id="xref-ref-1-2" class="xref-bibr" href="http://science.sciencemag.org/content/275/5301/737.5#ref-1">1</a>), humans may indirectly increase N inputs to terrestrial ecosystms by increasing atmospheric CO2 concentrations. The interaction between CO2 and N enrichment, as well as shifts in plant species, will likely influence future C storage by the terrestrial biosphere</p>
Hungate BA, Lund CP, Pearson HL, CHAPIN III FS (1997) Elevated CO2 and nutrient addition after soil N cycling and N trace gas fluxes with early season wet-up in a California annual grassland. Biogeochemistry 37(2): 89-109.
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Download .PDFWe examined the effects of growth carbon dioxide (CO<sub>2</sub>)concentration and soil nutrient availability on nitrogen (N)transformations and N trace gas fluxes in California grasslandmicrocosms during early-season wet-up, a time when rates of Ntransformation and N trace gas flux are high. After plant senescenceand summer drought, we simulated the first fall rains and examined Ncycling. Growth at elevated CO<sub>2</sub> increased root productionand root carbon:nitrogen ratio. Under nutrient enrichment, elevatedCO<sub>2</sub> increased microbial N immobilization during wet-up,leading to a 43% reduction in gross nitrification anda 55% reduction in NO emission from soil. ElevatedCO<sub>2</sub> increased microbial N immobilization at ambientnutrients, but did not alter nitrification or NO emission. ElevatedCO<sub>2</sub> did not alter soil emission of N<sub>2</sub>O ateither nutrient level. Addition of NPK fertilizer (1:1:1) stimulatedN mineralization and nitrification, leading to increased N<sub>2</sub>Oand NO emission from soil. The results of our study support a mechanisticmodel in which elevated CO<sub>2</sub> alters soil N cycling and NOemission: increased root production and increased C:N ratio in elevatedCO<sub>2</sub> stimulate N immobilization, thereby decreasingnitrification and associated NO emission when nutrients are abundant.This model is consistent with our basic understanding of how C availabilityinfluences soil N cycling and thus may apply to many terrestrial ecosystems.
Reynolds HL, Hungate BA, Chapin III FS, D'Antonio CM (1997) Soil heterogeneity and plant competition in anannual grassland. Ecology 78(7): 2076-2090.
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Read PublicationVariation in competitive ability due to variation in soil characteristics is one possible mechanism allowing the local coexistence of plant species. We measured soil water, depth, and nitrogen pools and fluxes in distinct patches of three serpentine grassland species to determine whether soil heterogeneity existed and was correlated with plant species abundance. Through experimental manipulation of species’ abundances, we also examined the relative importance of inherent site characteristics vs. plant species’ effects in generating heterogeneity in the measured soil characteristics; and measured species’ competitive abilities in different patch types. The three common grassland annuals, <em>Calycadenia multiglandulosum, Plantago erecta,</em> and <em>Lasthenia californica,</em> were segregated with respect to the measured soil characteristics. Differences in soil water, soil depth, soil microbial nitrogen, and soil carbon to nitrogen ratio were due to inherent site characteristics, while differences in nitrate availability were strongly affected by the identity of the species currently growing in a soil patch. Furthermore, all species performed significantly better against one other species in the patch type where they are normally most abundant. These results demonstrate that species diversity within this grassland contributes to soil heterogeneity and suggest that soil heterogeneity could contribute to the coexistence of these species.
1996
Hungate BA, Canadell J, Chapin FS (1996) Plant species mediate changes in soil microbial N in response to elevated CO2. Ecology 2505-2515.
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Read PublicationThe effect of elevated CO2 on plant-microbial interactions and nitrogen (N) cycling is critical to predicting plant growth responses to elevated CO2, because plant growth is often N-limited. We investigated whether the effects of elevated CO2 on plant-microbial N dynamics differed among six annual plant species: three European grasses that have invaded California grasslands, and one grass and two forbs native to California serpentine grassland. Elevated CO2 altered plant N pools and '5NH4+ uptake, but the direction and magnitude of the changes were species dependent. The introduced grasses showed increased plant N pools and '5NH4+ uptake, whereas the native species showed smaller increases or even decreases in plant N pools and '5NH4+ uptake. Under nutrient enrichment, soil microbial N and '5NH4+ uptake differed among soils with different plant species, but they were not affected by elevated CO2. At low nutrients, elevated CO2 altered soil microbial N and '5NH4+ uptake, but the direction and magnitude of the changes were species dependent. The changes in soil microbial N were positively correlated with changes in the plant N pool, suggesting that there was no trade-off in N uptake between plants and microbes. These results also suggest that plant species composition will partly determine the direction of changes in soil N cycling in response to elevated CO2.
1995
Hungate BA, Jackson RB, Field CB, Chapin III FS (1995) Detecting changes in soil carbon in CO2 enrichment experiments. Plant and Soil 187(2): 135-145.
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Read PublicationAfter four growing seasons, elevated CO<sub>2</sub> did not significantly alter surface soil C pools in two intact annual grasslands. However, soil C pools in these systems are large compared to the likely changes caused by elevated CO<sub>2</sub>. We calculated statistical power to detect changes in soil C, using an approach applicable to all elevated CO<sub>2</sub> experiments. The distinctive isotopic signature of the fossil-fuel-derived CO<sub>2</sub> added to the elevated CO<sub>2</sub> treatment provides a C tracer to determine the rate of incorporation of newly-fixed C into soil. This rate constrains the size of the possible effect of eievated CO<sub>2</sub> on soil C. Even after four years of treatment, statistical power to detect plausible changes in soil C under elevated CO<sub>2</sub> is quite low. Analysis of other elevated CO<sub>2</sub> experiments in the literature indicates that either CO<sub>2</sub> does not affect soil C content, or that reported CO<sub>2</sub> effects on soil C are too large to be a simple consequence of increased plant carbon inputs, suggesting that other mechanisms are involved, or that the differences are due to chance. Determining the effects of elevated CO<sub>2</sub> on total soil C and long-term C storage requires more powerful experimental techniques or experiments of longer duration.
Mack MC, Syndonia Bret-Harte M, Hollingsworth TN, Jandt RR, Schuur EAG, Shaver GR, Verbyla DL () Carbon loss from an unprecedented Arctic tundra wildfire. .
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Read PublicationArctic tundra soils store large amounts of carbon (C) in organic soil layers hundreds to thousands of years old that insulate, and in some cases maintain, permafrost soilsl,2. Fire has been largely absent from most of this biome since the early Holocene epoch3, but its frequency and extent are increasing, probably in response to climate warming4. The effect of fires on the C balance of tundra landscapes, however, remains largely unknown. The Anaktuvuk River fire in 2007 burned 1,039 square kilometres of Alaska's Arctic slope, making it the largest fire on record for the tundra biome and doubling the cumulative area burned since 1950 (ref. 5). Here we report that tundra ecosystems lost 2,016 ± 435 g C m -2 in the fire, an amount two orders of magnitude larger than annual net C exchange in undisturbed tundra6• Sixty per cent of this C loss was from soil organic matter, and radiocarbon dating of residual soil layers revealed that the maximum age of soil C lost was 50 years. Scaled to the entire burned area, the fire released approximately 2.1 teragrams of C to the atmosphere, an amount similar in magnitude to the annual net C sink for the entire Arctic tundra biome averaged over the last quarter of the twentieth century". The magnitude of ecosystem C lost by fire, relative to both ecosystem and biome-scale fluxes, demonstrates that a climate-driven increase in tundra fire disturbance may represent a positive feedback, potentially offsetting Arctic greeningB and influencing the net C balance of the tundra biome.