Effect of winter fire on primary productivity and nutrient concentration of a dry tropical savanna

Burning increased the mean annual canopy and belowground biomass of a dry tropical savanna by 40% and 12%, respectively, while littermass was reduced by 85% in comparison to control savanna. Mean annual aboveground and belowground net primary production were 471 and 631 g m^-2 in control, and 584 and 688 g m^-2 in burned savanna, respectively. Fire caused an increase in mean aboveground net production of 24% and in belowground net production of 9%.Concentration of carbon, nitrogen and phosphorus in vegetation of unburned plots ranged between 34.01–38.59%, 0.85–1.53% and 0.04–0.11% and in soil from 0.95–1%, 0.011–0.13% and 0.017–0.02%, respectively. Fire increased the mean concentrations of N and P by 16% and 42% in vegetation and 18.18% and 17.65% in soil, respectively. Thus winter fire can be an important tool for the management of dry tropical savanna with respect to biomass production and nutritive quality.     

Experimental fire increases soil carbon dioxide efflux in a grassland long‐term multifactor global change experiment

Numerous studies have demonstrated that soil respiration rates increase under experimental warming, although the long‐term, multiyear dynamics of this feedback are not well constrained. Less is known about the effects of single, punctuated events in combination with other longer‐duration anthropogenic influences on the dynamics of soil carbon (C) loss. In 2012 and 2013, we assessed the effects of decadal‐scale anthropogenic global change – warming, increased nitrogen (N) deposition, elevated carbon dioxide (CO₂), and increased precipitation – on soil respiration rates in an annual‐dominated Mediterranean grassland. We also investigated how controlled fire and an artificial wet‐up event, in combination with exposure to the longer‐duration anthropogenic global change factors, influenced the dynamics of C cycling in this system. Decade‐duration surface soil warming (1–2 °C) had no effect on soil respiration rates, while +N addition and elevated CO₂ concentrations increased growing‐season soil CO₂ efflux rates by increasing annual aboveground net primary production (NPP) and belowground fine root production, respectively. Low‐intensity experimental fire significantly elevated soil CO₂ efflux rates in the next growing season. Based on mixed‐effects modeling and structural equation modeling, low‐intensity fire increased growing‐season soil respiration rates through a combination of three mechanisms: large increases in soil temperature (3–5 °C), significant increases in fine root production, and elevated aboveground NPP. Our study shows that in ecosystems where soil respiration has acclimated to moderate warming, further increases in soil temperature can stimulate greater soil CO₂ efflux. We also demonstrate that punctuated short‐duration events such as fire can influence soil C dynamics with implications for both the parameterization of earth system models (ESMs) and the implementation of climate change mitigation policies that involve land‐sector C accounting.     

Interactive Effects of Time, CO2, N, and Diversity on Total Belowground Carbon Allocation and Ecosystem Carbon Storage in a Grassland Community

Predicting if ecosystems will mitigate or exacerbate rising CO₂ requires understanding how elevated CO₂ will interact with coincident changes in diversity and nitrogen (N) availability to affect ecosystem carbon (C) storage. Yet achieving such understanding has been hampered by the difficulty of quantifying belowground C pools and fluxes. Thus, we used mass balance calculations to quantify the effects of diversity, CO₂, and N on both the total amount of C allocated belowground by plants (total belowground C allocation, TBCA) and ecosystem C storage in a periodically burned, 8-year Minnesota grassland biodiversity, CO₂, and N experiment (BioCON). Annual TBCA increased in response to elevated CO₂, enriched N, and increasing diversity. TBCA was positively related to standing root biomass. After removing the influence of root biomass, the effect of elevated CO₂ remained positive, suggesting additional drivers of TBCA apart from those that maintain high root biomass. Removing root biomass effects resulted in the effects of N and diversity becoming neutral or negative (depending on year), suggesting that the positive effects of diversity and N on TBCA were related to treatment-driven differences in root biomass. Greater litter production in high diversity, elevated CO₂, and enhanced N treatments increased annual ecosystem C loss in fire years and C gain in non-fire years, resulting in overall neutral C storage rates. Our results suggest that frequently burned grasslands are unlikely to exhibit enhanced C sequestration with increasing atmospheric CO₂ levels or N deposition.     

Nitrogen and phosphorus dynamics of a tallgrass prairie ecosystem exposed to elevated carbon dioxide

A tallgrass prairie ecosystem was exposed to ambient and twice-ambient CO₂ concentrations in open-top chambers and compared to unchambered ambient CO₂ during the entire growing season from 1989 through 1991. Dominant species were Andropogon gerardii (C₄), A. scoparius (C₄), Sorghastrum nutans (C₄) and Poa pratensis (C₃). Nitrogen and phosphorus concentrations in A. gerardii, P. pratensis and dicotyledonous herbs above ground biomass were estimated by periodic sampling throughout the growing season in 1989 and 1990. In 1991, N and P concentrations in peak biomass were estimated by an early August harvest. N and P concentrations in root production as a function of treatment were estimated using root ingrowth bags that remained in place throughout the growing season. Total N and P in above- and belowground biomass were calculated as products of concentration and peak biomass by species groups. N concentration in A. gerardii and dicotyledonous herb aboveground biomass was lower and total N higher in elevated CO₂ plots than in ambient CO₂ plots. N concentration in P. pratensis aboveground biomass was lower in elevated CO₂ plots than in ambient, but total N did not differ among treatments in 2 out of 3 years. In 1990, N concentration in root ingrowth bag biomass was lower and total N greater in elevated CO₂ than in ambient CO₂ plots. Root ingrowth bag biomass N concentration did not differ among treatments in 1991, but total N was greater in elevated CO₂ plots than in ambient CO₂ plots. P concentration was lower under elevated CO₂ compared to ambient in 1989, but did not differ substantially among treatments in 1990 or 1991. In all years, total P in aboveground A. gerardii and root ingrowth bag biomass was greater under elevated CO₂ than ambient. P concentration and total P in P. pratensis was similar among treatments.     

Nitrate supply and plant development influence nitrogen uptake and allocation under elevated CO<Subscript>2</Subscript> in durum wheat grown hydroponically

Growth in elevated CO₂ often leads to decreased plant nitrogen contents and down-regulation of photosynthetic capacity. Here, we investigated whether elevated CO₂ limits nitrogen uptake when nutrient movement to roots is unrestricted, and the dependence of this limitation on nitrogen supply and plant development in durum wheat (Triticum durum Desf.). Plants were grown hydroponically at two N supplies and ambient and elevated CO₂ concentrations. Elevated CO₂ decreased nitrate uptake per unit root mass with low N supply at early grain filling, but not at anthesis. This decrease was not associated with higher nitrate or amino acid, or lower non-structural carbohydrate contents in roots. At anthesis, elevated CO₂ decreased the nitrogen content of roots with both levels of N and that of aboveground organs with high N. With low N, elevated CO₂ increased N allocation to aboveground plant organs and nitrogen concentration per unit flag leaf area at anthesis, and per unit aboveground dry mass at both growth stages. The results from the hydroponic experiment suggest that elevated CO₂ restricts nitrate uptake late in development, high N supply overriding this restriction. Increased nitrogen allocation to young leaves at low N supply could alleviate photosynthetic acclimation to elevated CO₂.     

Plant Nitrogen Dynamics in Shortgrass Steppe under Elevated Atmospheric Carbon Dioxide

The direct and indirect effects of increasing levels of atmospheric carbon dioxide (CO₂) on plant nitrogen (N) content were studied in a shortgrass steppe ecosystem in northeastern Colorado, USA. Beginning in 1997 nine experimental plots were established: three open-top chambers with ambient CO₂ levels (approximately 365 mol mol^–1), three open-top chambers with twice-ambient CO₂ levels (approximately 720 mol mol^–1), and three unchambered control plots. After 3 years of growing-season CO₂ treatment, the aboveground N concentration of plants grown under elevated atmospheric CO₂ decreased, and the carbon–nitrogen (C:N) ratio increased. At the same time, increased aboveground biomass production under elevated atmospheric CO₂ conditions increased the net transfer of N out of the soil of elevated-CO₂ plots. Aboveground biomass production after simulated herbivory was also greater under elevated CO₂ compared to ambient CO₂. Surprisingly, no significant changes in belowground plant tissue N content were detected in response to elevated CO₂. Measurements of individual species at peak standing phytomass showed significant effects of CO₂ treatment on aboveground plant tissue N concentration and significant differences between species in N concentration, suggesting that changes in species composition under elevated CO₂ will contribute to overall changes in nutrient cycling. Changes in plant N content, driven by changes in aboveground plant N concentration, could have important consequences for biogeochemical cycling rates and the long-term productivity of the shortgrass steppe as atmospheric CO₂ concentrations increase.     

Biomass production in a nitrogen-fertilized,tallgrass prairie ecosystem exposed to ambient and elevated levels of CO2

Increased biomass production in terrestrial ecosystems with elevated atmospheric CO₂ may be constrained by nutrient limitations as a result of increased requirement or reduced availability caused by reduced turnover rates of nutrients. To determine the short-term impact of nitrogen (N) fertilization on plant biomass production under elevated CO₂, we compared the response of N-fertilized tallgrass prairie at ambient and twice-ambient CO₂ levels over a 2-year period. Native tallgrass prairie plots (4.5 m diameter) were exposed continuously (24 h) to ambient and twice-ambient CO₂ from 1 April to 26 October. We compared our results to an unfertilized companion experiment on the same research site. Above- and belowground biomass production and leaf area of fertilized plots were greater with elevated than ambient CO₂ in both years. The increase in biomass at high CO₂ occurred mainly aboveground in 1991, a dry year, and belowground in 1990, a wet year. Nitrogen concentration was lower in plants exposed to elevated CO₂, but total standing crop N was greater at high CO₂. Increased root biomass under elevated CO₂ apparently increased N uptake. The biomass production response to elevated CO₂ was much greater on N-fertilized than unfertilized prairie, particularly in the dry year. We conclude that biomass production response to elevated CO₂ was suppressed by N limitation in years with below-normal precipitation. Reduced N concentration in above- and belowground biomass could slow microbial degradation of soil organic matter and surface litter, thereby exacerbating N limitation in the long term.     

Sustained effects of atmospheric [CO2] and nitrogen availability on forest soil CO2 efflux

Soil CO₂ efflux (F_soil) is the largest source of carbon from forests and reflects primary productivity as well as how carbon is allocated within forest ecosystems. Through early stages of stand development, both elevated [CO₂] and availability of soil nitrogen (N; sum of mineralization, deposition, and fixation) have been shown to increase gross primary productivity, but the long‐term effects of these factors on F_soil are less clear. Expanding on previous studies at the Duke Free‐Air CO₂ Enrichment (FACE) site, we quantified the effects of elevated [CO₂] and N fertilization on F_soil using daily measurements from automated chambers over 10 years. Consistent with previous results, compared to ambient unfertilized plots, annual F_soil increased under elevated [CO₂] (ca. 17%) and decreased with N (ca. 21%). N fertilization under elevated [CO₂] reduced F_soil to values similar to untreated plots. Over the study period, base respiration rates increased with leaf productivity, but declined after productivity saturated. Despite treatment‐induced differences in aboveground biomass, soil temperature and water content were similar among treatments. Interannually, low soil water content decreased annual F_soil from potential values – estimated based on temperature alone assuming nonlimiting soil water content – by ca. 0.7% per 1.0% reduction in relative extractable water. This effect was only slightly ameliorated by elevated [CO₂]. Variability in soil N availability among plots accounted for the spatial variability in F_soil, showing a decrease of ca. 114 g C m^?2 yr^?1 per 1 g m^?2 increase in soil N availability, with consistently higher F_soil in elevated [CO₂] plots ca. 127 g C per 100 ppm [CO₂] over the +200 ppm enrichment. Altogether, reflecting increased belowground carbon partitioning in response to greater plant nutritional needs, the effects of elevated [CO₂] and N fertilization on F_soil in this stand are sustained beyond the early stages of stand development and through stabilization of annual foliage production.     

Variation in Aboveground Cover Influences Soil Nitrogen Availability at Fine Spatial Scales Following Severe Fire in Subalpine Conifer Forests

Following fire, fine-scale variation in early successional vegetation and soil nutrients may influence development of ecosystem structure and function. We studied conifer forests burned by stand-replacing wildfire in Greater Yellowstone (Wyoming, USA) to address two questions: (1) How do the variability and spatial structure of aboveground cover and soil nitrogen availability change during the first 4 years following stand-replacing fire? (2) At fine scales (2–20 m), are postfire soil inorganic N pools and fluxes related to aboveground cover? Aboveground cover, soil N pools, and annual net N transformations were measured from 2001 to 2004 using a spatially explicit sampling design in four 0.25-ha plots that burned during summer 2000. Within-stand variability (coefficient of variation) in postfire live vegetative cover declined with time since fire, whereas variability in bare mineral soil, charred litter and fresh litter was greatest 2-3 years postfire. The soil nitrate pool was more variable than the soil ammonium pool, but annual net nitrification was less variable than annual net N mineralization. Spatial structure (quantified by semivariograms) was observed in some aboveground cover variables (for example, graminoids and fresh litter), but there was little spatial structure in soil N variables and no obvious congruence in spatial scales of autocorrelation for soil N and aboveground cover. Significant Spearman correlations (at the sample point) indicated that aboveground cover and soil N were coupled following severe fire, and the dominant influence was from aboveground cover to soil N, rather than from soil N to vegetation. Initial patterns of fire severity and re-vegetation contributed to fine-scale heterogeneity in soil N availability for at least 4 years after severe fire.     

Growth of Eastern Cottonwoods (Populus deltoides) in elevated [CO2] stimulates stand-level respiration and rhizodeposition of carbohydrates, accelerates soil nutrient depletion, yet stimulates above- and belowground biomass production

We took advantage of the distinctive system‐level measurement capabilities of the Biosphere 2 Laboratory (B2L) to examine the effects of prolonged exposure to elevated [CO₂] on carbon flux dynamics, above‐ and belowground biomass changes, and soil carbon and nutrient capital in plantation forest stands over 4 years. Annually coppiced stands of eastern cottonwoods (Populus deltoides) were grown under ambient (400 ppm) and two levels of elevated (800 and 1200 ppm) atmospheric [CO₂] in carbon and N‐replete soils of the Intensive Forestry Mesocosm in the B2L. The large semiclosed space of B2L uniquely enabled precise CO₂ exchange measurements at the near ecosystem scale. Highly controllable climatic conditions within B2L also allowed for reproducible examination of CO₂ exchange under different scales in space and time. Elevated [CO₂] significantly stimulated whole‐system maximum net CO₂ influx by an average of 21% and 83% in years 3 and 4 of the experiment. Over the 4‐year experiment, cumulative belowground, foliar, and total aboveground biomass increased in both elevated [CO₂] treatments. After 2 years of growth at elevated [CO₂], early season stand respiration was decoupled from CO₂ influx aboveground, presumably because of accelerated fine root production from stored carbohydrates in the coppiced system prior to canopy development and to the increased soil carbohydrate status under elevated [CO₂] treatments. Soil respiration was stimulated by elevated [CO₂] whether measured at the system level in the undisturbed soil block, by soil collars in situ, or by substrate‐induced respiration in vitro. Elevated [CO₂] accelerated depletion of soil nutrients, phosphorus, calcium and potassium, after 3 years of growth, litter removal, and coppicing, especially in the upper soil profile, although total N showed no change. Enhancement of above‐ and belowground biomass production by elevated [CO₂] accelerated carbon cycling through the coppiced system and did not sequester additional carbon in the soil.     

Annual burning of a tallgrass prairie inhibits C and N cycling in soil,increasing recalcitrant pyrogenic organic matter storage while reducing N availability

Grassland ecosystems store an estimated 30% of the world's total soil C and are frequently disturbed by wildfires or fire management. Aboveground litter decomposition is one of the main processes that form soil organic matter (SOM). However, during a fire biomass is removed or partially combusted and litter inputs to the soil are substituted with inputs of pyrogenic organic matter (py‐OM). Py‐OM accounts for a more recalcitrant plant input to SOM than fresh litter, and the historical frequency of burning may alter C and N retention of both fresh litter and py‐OM inputs to the soil. We compared the fate of these two forms of plant material by incubating ¹³C‐ and ¹⁵N‐labeled Andropogon gerardii litter and py‐OM at both an annually burned and an infrequently burned tallgrass prairie site for 11 months. We traced litter and py‐OM C and N into uncomplexed and organo‐mineral SOM fractions and CO₂ fluxes and determined how fire history affects the fate of these two forms of aboveground biomass. Evidence from CO₂ fluxes and SOM C:N ratios indicates that the litter was microbially transformed during decomposition while, besides an initial labile fraction, py‐OM added to SOM largely untransformed by soil microbes. Additionally, at the N‐limited annually burned site, litter N was tightly conserved. Together, these results demonstrate how, although py‐OM may contribute to C and N sequestration in the soil due to its resistance to microbial degradation, a long history of annual removal of fresh litter and input of py‐OM infers N limitation due to the inhibition of microbial decomposition of aboveground plant inputs to the soil. These results provide new insight into how fire may impact plant inputs to the soil, and the effects of py‐OM on SOM formation and ecosystem C and N cycling.     

Controls of nitrogen limitation in tallgrass prairie

Summary The relationship between fire frequency and N limitation to foliage production in tallgrass prairie was studied with a series of fire and N addition experiments. Results indicated that fire history affected the magnitude of the vegetation response to fire and to N additions. Sites not burned for over 15 years averaged only a 9% increase in foliage biomass in response to N enrichment. In contrast, foliage production increased an average of 68% in response to N additions on annually burned sites, while infrequently burned sites, burned in the year of the study, averaged a 45% increase. These findings are consistent with reports indicating that reduced plant growth on unburned prairie is due to shading and lower soil temperatures, while foliage production on frequently burned areas is constrained by N availability. Infrequent burning of unfertilized prairie therefore results in a maximum production response in the year of burning relative to either annually burned or long-term unburned sites.Foliage biomass of tallgrass prairie is dominated by C₄ grasses; however, forb species exhibited stronger production responses to nitrogen additions than did the grasses. After four years of annual N additions, forb biomass exceeded that of grass biomass on unburned plots, and grasses exhibited a negative response to fertilizer, probably due to competition from the forbs. The dominant C₄ grasses may out-compete forbs under frequent fire conditions not only because they are better adapted to direct effects of burning, but because they can grow better under low available N regimes created by frequent fire.     

Biomass production and species composition change in a tallgrass prairie ecosystem after long-term exposure to elevated atmospheric CO2

To determine the long-term impact of elevated CO₂ on primary production of native tallgrass prairie, we compared the responses of tallgrass prairie at ambient and twice-ambient atmospheric CO₂ levels over an 8-year period. Plots in open-top chambers (4.5 m diameter) were exposed continuously (24 h) to ambient and elevated CO₂ from early April to late October each year. Unchambered plots were monitored also. Above-ground peak biomass was determined by clipping each year in early August, and root growth was estimated by harvesting roots from root ingrowth bags. Plant community composition was censused each year in early June. In the last 2 years of the study, subplots were clipped on 1 June or 1 July, and regrowth was harvested on 1 October. Volumetric soil water content of the 0–100 cm soil layer was determined using neutron scattering, and was generally higher in elevated CO₂ plots than ambient. Peak above-ground biomass was greater on elevated CO₂ plots than ambient CO₂ plots with or without chambers during years with significant plant water stress. Above-ground regrowth biomass was greater under elevated CO₂ than under ambient CO₂ in a year with late-season water stress, but did not differ in a wetter year. Root ingrowth biomass was also greater in elevated CO₂ plots than ambient CO₂ plots when water stress occurred during the growing season. The basal cover and relative amount of warm-season perennial grasses (C4) in the stand changed little during the 8-year period, but basal cover and relative amount of cool-season perennial grasses (C3) in the stand declined in the elevated CO₂ plots and in ambient CO₂ plots with chambers. Forbs (C3) and members of the Cyperaceae (C3) increased in basal cover and relative amount in the stand at elevated compared to ambient CO₂. Greater biomass production under elevated CO₂ in C4-dominated grasslands may lead to a greater carbon sequestration by those ecosystems and reduce peak atmospheric CO₂ concentrations in the future.     

Effects of elevated CO2, temperature and N fertilization on nitrogen fluxes in a temperate grassland ecosystem

The soil nitrogen cycle was investigated in a pre‐established Lolium perenne sward on a loamy soil and exposed to ambient and elevated atmospheric CO₂ concentrations (350 and 700 μL L^?1) and, at elevated [CO₂], to a 3 °C temperature increase. At two levels of mineral nitrogen supply, N– (150 kgN ha^?1 y^?1) and N+ (533 kgN ha^?1 y^?1), ¹⁵N‐labelled ammonium nitrate was supplied in split applications over a 2.5‐y period. The recovery of the labelled fertilizer N was measured in the harvests, in the stubble and roots, in the macro‐organic matter fractions above 200 μm in size (MOM) and in the aggregated organic matter below 200 μM (AOM). Elevated [CO₂] reduced the total amount of N harvested in the clipped parts of the sward. The harvested N derived from soil was reduced to a greater extent than that derived from fertilizer. At both N supplies, elevated [CO₂] modified the allocation of the fertilizer N in the sward, in favour of the stubble and roots and significantly increased the recovery of fertilizer N in the soil macro‐organic matter fractions. The increase of fertilizer N immobilization in the MOM was associated with a decline of fertilizer N uptake by the grass sward, which supported the hypothesis of a negative feedback of elevated [CO₂] on the sward N yield and uptake. Similar and even more pronounced effects were observed for the native N mineralized in the soil. At N–, a greater part of the fertilizer N organized in the root phytomass resulted in an underestimation of N immobilized in dead roots and, in turn, an underestimation of N immobilization in the MOM. The 3 °C temperature increase alleviated the [CO₂] effect throughout much of the N cycle, increasing soil N mineralization, N derived from soil in the harvests, and the partitioning of the assimilated fertilizer N to shoots. In conclusion, at ambient temperature, the N cycle was slowed down under elevated [CO₂], which restricted the increase in the aboveground production of the grass sward, and apparently contributed to the sequestration of carbon belowground. In contrast, a temperature increase under elevated [CO₂] stimulated the soil nitrogen cycle, improved the N nutrition of the sward and restricted the magnitude of the soil C sequestration.     

Species-specific responses of plant communities to altered carbon and nutrient availability

In a field microcosm experiment, species‐specific responses of aboveground biomass of two California annual grassland communities to elevated CO₂ and nutrient availability were investigated. One community grows on shallow, nutrient‐poor serpentine‐derived soil whereas the other occurs on deeper, modestly fertile sandstone/greenstone‐derived substrate. In most species, CO₂ effects did not appear until late in the growing season, probably because the elevated CO₂ increased water‐use‐efficiency easing, the onset of the summer drought. Responses of aboveground biomass to elevated CO₂ differed depending on nutrient availability. Similarly, biomass responses to nutrient treatments differed depending on the CO₂ status. For the majority of the species, production increased most under elevated CO₂ with added nutrients (N,P,K, and micro nutrients). Some species were losers under conditions that increased overall community production, including Bromus hordeaceus in the serpentine community (negative biomass response under elevated CO₂) and Lotus wrangelianus in both communities (negative biomass response with added nitrogen). Treatment and competitive effects on species‐specific biomass varied in both magnitude and direction, especially in the serpentine community, significantly affecting community structure. Individual resource environments are likely to be affected by neighbouring plants, and these competitive interactions complicate predictions of species' responses to elevated CO₂.     

Decomposition of soybean grown under elevated concentrations of CO2 and O3

A critical global climate change issue is how increasing concentrations of atmospheric CO₂ and ground‐level O₃ will affect agricultural productivity. This includes effects on decomposition of residues left in the field and availability of mineral nutrients to subsequent crops. To address questions about decomposition processes, a 2‐year experiment was conducted to determine the chemistry and decomposition rate of aboveground residues of soybean (Glycine max (L.) Merr.) grown under reciprocal combinations of low and high concentrations of CO₂ and O₃ in open‐top field chambers. The CO₂ treatments were ambient (370 μmol mol^?1) and elevated (714 μmol mol^?1) levels (daytime 12 h averages). Ozone treatments were charcoal‐filtered air (21 nmol mol^?1) and nonfiltered air plus 1.5 times ambient O₃ (74 nmol mol^?1) 12 h day^?1. Elevated CO₂ increased aboveground postharvest residue production by 28–56% while elevated O₃ suppressed it by 15–46%. In combination, inhibitory effects of added O₃ on biomass production were largely negated by elevated CO₂. Plant residue chemistry was generally unaffected by elevated CO₂, except for an increase in leaf residue lignin concentration. Leaf residues from the elevated O₃ treatments had lower concentrations of nonstructural carbohydrates, but higher N, fiber, and lignin levels. Chemical composition of petiole, stem, and pod husk residues was only marginally affected by the elevated gas treatments. Treatment effects on plant biomass production, however, influenced the content of chemical constituents on an areal basis. Elevated CO₂ increased the mass per square meter of nonstructural carbohydrates, phenolics, N, cellulose, and lignin by 24–46%. Elevated O₃ decreased the mass per square meter of these constituents by 30–48%, while elevated CO₂ largely ameliorated the added O₃ effect. Carbon mineralization rates of component residues from the elevated gas treatments were not significantly different from the control. However, N immobilization increased in soils containing petiole and stem residues from the elevated CO₂, O₃, and combined gas treatments. Mass loss of decomposing leaf residue from the added O₃ and combined gas treatments was 48% less than the control treatment after 20 weeks, while differences in decomposition of petiole, stem, and husk residues among treatments were minor. Decreased decomposition of leaf residues was correlated with lower starch and higher lignin levels. However, leaf residues only comprised about 20% of the total residue biomass assayed so treatment effects on mass loss of total aboveground residues were relatively small. The primary influence of elevated atmospheric CO₂ and O₃ concentrations on decomposition processes is apt to arise from effects on residue mass input, which is increased by elevated CO₂ and suppressed by O₃.     

Elevated atmospheric CO2 in open top chambers increases net nitrification and potential denitrification

The control of soil nitrogen (N) availability under elevated atmospheric CO₂ is central to predicting changes in ecosystem carbon (C) storage and primary productivity. The effects of elevated CO₂ on belowground processes have so far attracted limited research and they are assumed to be controlled by indirect effects through changes in plant physiology and chemistry. In this study, we investigated the effects of a 4‐year exposure to elevated CO₂ (ambient + 400 µmol mol^?1) in open top chambers under Scots pine (Pinus sylvestris L) seedlings on soil microbial processes of nitrification and denitrification. Potential denitrification (DP) and potential N₂O emissions were significantly higher in soils from the elevated CO₂ treatment, probably regulated indirectly by the changes in soil conditions (increased pH, C availability and NO₃^– production). Net N mineralization was mainly accounted for by nitrate production. Nitrate production was significantly larger for soil from the elevated CO₂ treatment in the field when incubated in the laboratory under elevated CO₂ (increase of 100%), but there was no effect when incubated under ambient CO₂. Net nitrate production of the soil originating from the ambient CO₂ treatment in the field was not influenced by laboratory incubation conditions. These results indicate that a direct effect of elevated atmospheric CO₂ on soil microbial processes might take place. We hypothesize that physiological adaptation or selection of nitrifiers could occur under elevated CO₂ through higher soil CO₂ concentrations. Alternatively, lower microbial NH₄ assimilation under elevated CO₂ might explain the higher net nitrification. We conclude that elevated atmospheric CO₂ has a major direct effect on the soil microbial processes of nitrification and denitrification despite generally higher soil CO₂ concentrations compared to atmospheric concentrations.     

Nitrification,denitrification, and related functional genes under elevated CO2: A meta-analysis in terrestrial ecosystems

Global change may have profound effects on soil nitrogen (N) cycling that can induce positive feedback to climate change through increased nitrous oxide (N₂O) emissions mediated by nitrification and denitrification. We conducted a meta-analysis of the effects of elevated CO₂ on nitrification and denitrification based on 879 observations from 58 publications and 46 independent elevated CO₂ experiments in terrestrial ecosystems. We investigated the effects of elevated CO₂ alone or combined with elevated temperature, increased precipitation, drought, and N addition. We assessed the response to elevated CO₂ of gross and potential nitrification, potential denitrification, and abundances of related functional genes (archaeal amoA, bacterial amoA, nirK, nirS, and nosZ). Elevated CO₂ increased potential nitrification (+28%) and the abundance of bacterial amoA functional gene (+62%) in cropland ecosystems. Elevated CO₂ increased potential denitrification when combined with N addition and higher precipitation (+116%). Elevated CO₂ also increased the abundance of nirK (+25%) and nirS (+27%) functional genes in terrestrial ecosystems and of nosZ (+32%) functional gene in cropland ecosystems. The increase in the abundance of nosZ under elevated CO₂ was larger at elevated temperature and high N (+62%). Four out of 14 two-way interactions tested between elevated CO₂ and elevated temperature, elevated CO₂ and increased precipitation, and elevated CO₂ and N addition were marginally significant and mostly synergistic. The effects of elevated CO₂ on potential nitrification and abundances of bacterial amoA and nirS functional genes increased with mean annual temperature and mean annual precipitation. Our meta-analysis thus suggests that warming and increased precipitation in large areas of the world could reinforce positive responses of nitrification and denitrification to elevated CO₂ and urges the need for more investigations in the tropical zone and on interactive effects among multiple global change factors, as we may largely underestimate the effects of global change on soil N₂O emissions.     

Short-term soil inorganic N pulse after experimental fire alters invasive and native annual plant production in a Mojave Desert shrubland

Post-fire changes in desert vegetation patterns are known, but the mechanisms are poorly understood. Theory suggests that pulse dynamics of resource availability confer advantages to invasive annual species, and that pulse timing can influence survival and competition among species. Precipitation patterns in the American Southwest are predicted to shift toward a drier climate, potentially altering post-fire resource availability and consequent vegetation dynamics. We quantified post-fire inorganic N dynamics and determined how annual plants respond to soil inorganic nitrogen variability following experimental fires in a Mojave Desert shrub community. Soil inorganic N, soil net N mineralization, and production of annual plants were measured beneath shrubs and in interspaces during 6 months following fire. Soil inorganic N pools in burned plots were up to 1 g m⁻² greater than unburned plots for several weeks and increased under shrubs (0.5–1.0 g m⁻²) more than interspaces (0.1–0.2 g m⁻²). Soil NO₃ ⁻−N (nitrate−N) increased more and persisted longer than soil NH₄ ⁺−N (ammonium−N). Laboratory incubations simulating low soil moisture conditions, and consistent with field moisture during the study, suggest that soil net ammonification and net nitrification were low and mostly unaffected by shrub canopy or burning. After late season rains, and where soil inorganic N pools were elevated after fire, productivity of the predominant invasive Schismus spp. increased and native annuals declined. Results suggest that increased N availability following wildfire can favor invasive annuals over natives. Whether the short-term success of invasive species following fire will direct long-term species composition changes remains to be seen, yet predicted changes in precipitation variability will likely interact with N cycling to affect invasive annual plant dominance following wildfire.     

Global change could amplify fire effects on soil greenhouse gas emissions

Background

Little is known about the combined impacts of global environmental changes and ecological disturbances on ecosystem functioning, even though such combined impacts might play critical roles in shaping ecosystem processes that can in turn feed back to climate change, such as soil emissions of greenhouse gases.

Methodology/Principal Findings

We took advantage of an accidental, low-severity wildfire that burned part of a long-term global change experiment to investigate the interactive effects of a fire disturbance and increases in CO₂ concentration, precipitation and nitrogen supply on soil nitrous oxide (N₂O) emissions in a grassland ecosystem. We examined the responses of soil N₂O emissions, as well as the responses of the two main microbial processes contributing to soil N₂O production – nitrification and denitrification – and of their main drivers. We show that the fire disturbance greatly increased soil N₂O emissions over a three-year period, and that elevated CO₂ and enhanced nitrogen supply amplified fire effects on soil N₂O emissions: emissions increased by a factor of two with fire alone and by a factor of six under the combined influence of fire, elevated CO₂ and nitrogen. We also provide evidence that this response was caused by increased microbial denitrification, resulting from increased soil moisture and soil carbon and nitrogen availability in the burned and fertilized plots.

Conclusions/Significance

Our results indicate that the combined effects of fire and global environmental changes can exceed their effects in isolation, thereby creating unexpected feedbacks to soil greenhouse gas emissions. These findings highlight the need to further explore the impacts of ecological disturbances on ecosystem functioning in the context of global change if we wish to be able to model future soil greenhouse gas emissions with greater confidence.