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2019
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Soil 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.
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Climate 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.
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Plant 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.
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Variation 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.
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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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Fertilization 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.
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Manual 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.
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Wildfire 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.
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Soil 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.
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Land 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.
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Wildfire 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.
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The 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.
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Tundra 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.
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The 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.
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Dynamic global vegetation models are key tools for interpreting and forecasting the responses of terrestrial ecosystems to climatic variation and other drivers. They estimate plant growth as the outcome of the supply of carbon through photosynthesis. However, growth is itself under direct control, and not simply controlled by the amount of available carbon. Therefore predictions by current photosynthesis-driven models of large increases in future vegetation biomass due to increasing concentrations of atmospheric CO2may be significant over-estimations. We describe how current understanding of wood formation can be used to reformulate global vegetation models, with potentially major implications for their behaviour.
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Dynamic global vegetation models are key tools for interpreting and forecasting the responses of terrestrial ecosystems to climatic variation and other drivers. They estimate plant growth as the outcome of the supply of carbon through photosynthesis. However, growth is itself under direct control, and not simply controlled by the amount of available carbon. Therefore predictions by current photosynthesis-driven models of large increases in future vegetation biomass due to increasing concentrations of atmospheric CO2may be significant over-estimations. We describe how current understanding of wood formation can be used to reformulate global vegetation models, with potentially major implications for their behaviour.
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It 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.
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Accurate 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.
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Reforestation 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.
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Forests 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.
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Plant 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.
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Determining 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.
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The 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.
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The 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.
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The 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.
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<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>
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Forest 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.
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Organic 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.
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The 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.
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Nutrient 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.
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Rice 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.
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Ongoing 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.
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Climate 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.
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Accurate 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.
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Warming 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.
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Shifts 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.
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Relationships 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.
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Global 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.
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An 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.
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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.
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Clarifying 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.
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How 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.
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As 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.
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Arctic 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.
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Organisms 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.
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Recent 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.
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Chinese 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.
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Ecologists 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).
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RNA 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.
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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.
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Higher 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.
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While 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.
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Evidence 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.
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Fire 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.
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Advancing 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.
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Microorganisms 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.
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Soil 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.
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It 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.
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Carbon 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.
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Monitoring 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.
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Digital 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.
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Direct 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.
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Temperature 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.
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Warming 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.
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Elevated 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.
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Temperature 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.
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Increasing 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.
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Boreal 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.
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Crop 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.
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One 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.
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Hydraulic 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.
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Piling 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.
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Carbon 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.
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Tree 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.
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Mulching 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.
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The 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.
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Soil fungal communities play a critical role in ecosystem carbon (C) and nitrogen (N) cycling. Although the effect of plant invasions on ecosystem C and N cycling is well established, its impact on soil fungal communities is not fully understood. The objective of this study was therefore to understand the variations in soil fungal communities as affected by plant invasion, and the mechanisms that drive these changes.
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In 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.
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Plant 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.
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Aim 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.
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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.
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Boreal 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.
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Boreal 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.
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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.
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Asymbiotic 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.
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Biological 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.
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Estimating 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.