Interactive Effects of Elevated CO2, Drought, and Warming on Plants

Adverse climate change attributed to elevated atmospheric carbon dioxide concentration (CO₂) and increased temperature components of global warming has been a central issue affecting economic and social development. Climate change, particularly global warming, imposes a severe impact on the terrestrial ecosystem. Elevated CO₂, drought, and high temperature have been extensively documented individually; however, relatively little is known about how plants respond to the interaction of these factors. To summarize current knowledge on the response of plants to global change factors, we focus on the interactive effects of CO₂ enrichment, warming, and drought on plant growth, carbon allocation, and photosynthesis. Stimulation due to elevated CO₂ might be suppressed under other negative climatic/environmental stresses such as drought, high temperature, and their combination. However, elevated CO₂ could alleviate deleterious effects of moderate drought via reducing stomatal conductance, altering leaf surface, and regulating gene expression. High CO₂ levels and rising temperatures may result in opposite responses in plant water use efficiency. Stimulation of plant growth due to elevated CO₂ for C₃ species occurs regardless of water conditions, but only under a water deficit for C₄ species. The positive effect of elevated CO₂ on C₄ species is derived mainly from the improved water status. Plant adaptive or maladaptive responses to multivariate environments are interactive; thus, researchers need to explore the ecological underpinnings involved in such responses to the multiple factors involved in climate change.     

Responses of Plant Community Composition and Biomass Production to Warming and Nitrogen Deposition in a Temperate Meadow Ecosystem

Climate change has profound influences on plant community composition and ecosystem functions. However, its effects on plant community composition and biomass production are not well understood. A four-year field experiment was conducted to examine the effects of warming, nitrogen (N) addition, and their interactions on plant community composition and biomass production in a temperate meadow ecosystem in northeast China. Experimental warming had no significant effect on plant species richness, evenness, and diversity, while N addition highly reduced the species richness and diversity. Warming tended to reduce the importance value of graminoid species but increased the value of forbs, while N addition had the opposite effect. Warming tended to increase the belowground biomass, but had an opposite tendency to decrease the aboveground biomass. The influences of warming on aboveground production were dependent upon precipitation. Experimental warming had little effect on aboveground biomass in the years with higher precipitation, but significantly suppressed aboveground biomass in dry years. Our results suggest that warming had indirect effects on plant production via its effect on the water availability. Nitrogen addition significantly increased above- and below-ground production, suggesting that N is one of the most important limiting factors determining plant productivity in the studied meadow steppe. Significant interactive effects of warming plus N addition on belowground biomass were also detected. Our observations revealed that environmental changes (warming and N deposition) play significant roles in regulating plant community composition and biomass production in temperate meadow steppe ecosystem in northeast China.     

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.     

Separating Drought Effects from Roof Artifacts on Ecosystem Processes in a Grassland Drought Experiment

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Given the predictions of increased drought probabilities under various climate change scenarios, there have been numerous experimental field studies simulating drought using transparent roofs in different ecosystems and regions. Such roofs may, however, have unknown side effects, called artifacts, on the measured variables potentially confounding the experimental results. A roofed control allows the quantification of potential artifacts, which is lacking in most experiments.

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We conducted a drought experiment in experimental grasslands to study artifacts of transparent roofs and the resulting effects of artifacts on ecosystems relative to drought on three response variables (aboveground biomass, litter decomposition and plant metabolite profiles). We established three drought treatments, using (1) transparent roofs to exclude rainfall, (2) an unroofed control treatment receiving natural rainfall and (3) a roofed control, nested in the drought treatment but with rain water reapplied according to ambient conditions.

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Roofs had a slight impact on air (+0.14°C during night) and soil temperatures (−0.45°C on warm days, +0.25°C on cold nights), while photosynthetically active radiation was decreased significantly (−16%). Aboveground plant community biomass was reduced in the drought treatment (−41%), but there was no significant difference between the roofed and unroofed control, i.e., there were no measurable roof artifact effects.

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Compared to the unroofed control, litter decomposition was decreased significantly both in the drought treatment (−26%) and in the roofed control treatment (−18%), suggesting artifact effects of the transparent roofs. Moreover, aboveground metabolite profiles in the model plant species Medicago x varia were different from the unroofed control in both the drought and roofed control treatments, and roof artifact effects were of comparable magnitude as drought effects.

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Our results stress the need for roofed control treatments when using transparent roofs for studying drought effects, because roofs can cause significant side effects.     

Effect of Sheep Urine Deposition on the Bacterial Community Structure in an Acidic Upland Grassland Soil

The effect of the addition of synthetic sheep urine (SSU) and plant species on the bacterial community composition of upland acidic grasslands was studied using a microcosm approach. Low, medium, and high concentrations of SSU were applied to pots containing plant species typical of both unimproved (Agrostis capillaris) and agriculturally improved (Lolium perenne) grasslands, and harvests were carried out 10 days and 50 days after the addition of SSU. SSU application significantly increased both soil pH (P < 0.005), with pH values ranging from pH 5.4 (zero SSU) to pH 6.4 (high SSU), and microbial activity (P < 0.005), with treatment with medium and high levels of SSU displaying significantly higher microbial activity (triphenylformazan dehydrogenase activity) than treatment of soil with zero or low concentrations of SSU. Microbial biomass, however, was not significantly altered by any of the SSU applications. Plant species alone had no effect on microbial biomass or activity. Bacterial community structure was profiled using bacterial automated ribosomal intergenic spacer analysis. Multidimensional scaling plots indicated that applications of high concentrations of SSU significantly altered the bacterial community composition in the presence of plant species but at different times: 10 days after application of high concentrations of SSU, the bacterial community composition of L. perenne-planted soils differed significantly from those of any other soils, whereas in the case of A. capillaris-planted soils, the bacterial community composition was different 50 days after treatment with high concentrations of SSU. Canonical correspondence analysis also highlighted the importance of interactions between SSU addition, plant species, and time in the bacterial community structure. This study has shown that the response of plants and bacterial communities to sheep urine deposition in grasslands is dependent on both the grass species present and the concentration of SSU applied, which may have important ecological consequences for agricultural grasslands.     

Complex Interrelationships Among Aboveground Biomass,Soil Chemical Properties,and Events Caused by Feral Goats and Their Eradication in a Grassland Ecosystem of an Island

This study examined the recovery, via biotic and abiotic pathways, of a grassland ecosystem after eradication of introduced exotic goats. We used path analyses to evaluate the relative strength of relationships among aboveground biomass, soil chemical properties (carbon, nitrogen, and phosphorus content; soil acidity), presence of nesting seabirds after goat eradication, extent of vegetation degraded by goats before their eradication, plant species composition after removal of goats, and topography. Models including the same variables with different paths were constructed using the Bayesian estimation method, and the best-fit models were constructed by comparing deviance information criterion values. Results of the path analyses demonstrated that vegetation degradation and soil erosion prior to goat eradication increased soil exchangeable acidity, which resulted in limitation of aboveground biomass. Seabird nesting after goat eradication increased the quantity of soil nutrients, possibly through inputs of feces, eggshells, and dead chicks or adults. The increase in nutrients was affected indirectly, via seabird nesting, by topography and vegetation type after goat eradication. The direct and indirect relationships demonstrated by our results suggest the existence of complex interrelationships during recovery of ecosystem function after eradication of exotic mammals.     

Effects of Elevated Carbon Dioxide on Plant Biomass Production and Competition in a Simulated Neutral Grassland Community

Using open-top chambers, four prominent species (Lolium perenne,Cynosurus cristatus, Holcus lanatusandAgrostis capillaris) ofIrish neutral grasslands were grown at ambient and elevated(700 µmol mol^-1) atmospheric CO₂for a period of 8 months.The effects of interspecific competition on plant responsesto CO₂enrichment were investigated by growing the species ina four-species mixture. The results indicate that the speciesdiffer in their ability to respond to elevated CO₂. CO₂-enrichmenthad the largest effect on the biomass production ofH. lanatus,but substantial stimulations in biomass production were alsofound for the other three species. The CO₂-stimulation of biomassproduction forH. lanatuswas accompanied by increased tillering.In addition, reductions in specific leaf area were found forall species. Exposure to elevated CO₂increased the communitybiomass of the four-species mixture. This increase can be mainlyattributed to a significant increase in the biomass ofH. lanatusatelevated CO₂. No statistically-significant changes in speciescomposition of community biomass were found. However,H. lanatusdidincrease its share of community biomass at each of the harvests,with the other three species, mainlyL. perenne, suffering lossesin their shares at elevated CO₂. The results show that: (1)the species varied in their response to elevated CO₂; and (2)species composition in natural plant communities is likely tochange at elevated CO₂, but these changes may occur rather slowly.Much longer periods of exposure to elevated atmospheric CO₂maybe required to permit detection of significant changes in speciescomposition.Copyright 1998 Annals of Botany Company Carbon dioxide (CO₂) enrichment, competition, Lolium perenne,Cynosurus cristatus, Holcus lanatus, Agrostis capillaris, biomass, specific leaf area, tillering.     

Warming and Elevated CO2 Interact to Drive Rapid Shifts in Marine Community Production

Predicting the outcome of future climate change requires an understanding of how alterations in multiple environmental factors manifest in natural communities and affect ecosystem functioning. We conducted an in situ, fully factorial field manipulation of CO₂ and temperature on a rocky shoreline in southeastern Alaska, USA. Warming strongly impacted functioning of tide pool systems within one month, with the rate of net community production (NCP) more than doubling in warmed pools under ambient CO₂ levels relative to initial NCP values. However, in pools with added CO₂, NCP was unaffected by warming. Productivity responses paralleled changes in the carbon-to-nitrogen ratio of a red alga, the most abundant primary producer species in the system, highlighting the direct link between physiology and ecosystem functioning. These observed changes in algal physiology and community productivity in response to our manipulations indicate the potential for natural systems to shift rapidly in response to changing climatic conditions and for multiple environmental factors to act antagonistically.     

Effects of Elevated Atmospheric CO2 on Soil Microbial Biomass, Activity, and Diversity in a Chaparral Ecosystem

This study reports the effects of long-term elevated atmospheric CO₂ on root production and microbial activity, biomass, and diversity in a chaparral ecosystem in southern California. The free air CO₂ enrichment (FACE) ring was located in a stand dominated by the woody shrub Adenostoma fasciculatum. Between 1995 and 2003, the FACE ring maintained an average daytime atmospheric CO₂ concentration of 550 ppm. During the last two years of operation, observations were made on soil cores collected from the FACE ring and adjacent areas of chaparral with ambient CO₂ levels. Root biomass roughly doubled in the FACE plot. Microbial biomass and activity were related to soil organic matter (OM) content, and so analysis of covariance was used to detect CO₂ effects while controlling for variation across the landscape. Extracellular enzymatic activity (cellulase and amylase) and microbial biomass C (chloroform fumigation-extraction) increased more rapidly with OM in the FACE plot than in controls, but glucose substrate-induced respiration (SIR) rates did not. The metabolic quotient (field respiration over potential respiration) was significantly higher in FACE samples, possibly indicating that microbial respiration was less C limited under high CO₂. The treatments also differed in the ratio of SIR to microbial biomass C, indicating a metabolic difference between the microbial communities. Bacterial diversity, described by 16S rRNA clone libraries, was unaffected by the CO₂ treatment, but fungal biomass was stimulated. Furthermore, fungal biomass was correlated with cellulase and amylase activities, indicating that fungi were responsible for the stimulation of enzymatic activity in the FACE treatment.     

Phenological Sensitivity of Early and Late Flowering Species Under Seasonal Warming and Altered Precipitation in a Seminatural Temperate Grassland Ecosystem

Shifts in flowering phenology of plants are indicators of climate change. The great majority of existing phenological studies refer solely to gradual warming. However, knowledge on how flowering phenology responds to changes in seasonal variation of warming and precipitation regimes is missing. We report the onset of 22 early (flowering before/within May) and 23 late flowering (flowering after May) species in response to manipulated seasonal warming (equal to + 1.2°C; last 100-year summer/winter warming), additional winter rainfall, and modified precipitation variability (including a 1000-year extreme drought event followed by heavy rainfall) over the growing season in two consecutive years for a species-rich temperate grassland ecosystem. The average onset of flowering (over 2 years) was significantly advanced 3.1 days by winter warming and 1.5 days by summer warming compared to control. Early flowering species responded to seasonal warming in both years, while late-flowering species responded in only 1 year to summer warming. The average onset of early flowering species was significantly advanced, 4.9 days by winter warming and 2.3 days by summer warming. Species-specific analysis showed that even within the early flowering community there were divergences. A positive correlation between plant height and shift in flowering onset was detected under winter warming (R² = 0.20, p = 0.005). The average onsets of early and late flowering community were affected by neither winter rain nor growing season precipitation variability. Seasonal differences in warming, and particularly winter warming, might alter community dynamics among early and late flowering species which can cause shifts in the seasonal performances of temperate ecosystems.     

Effects of Water Table Drawdown on Root Production and Aboveground Biomass in a Boreal Bog

We studied the effect of long-term water table drawdown on the vascular plant community in an ombrotrophic bog in central Finland by measuring aboveground biomass and belowground production (by in-growth cores) across plant functional groups including herbs, shrubs, and trees. We compared drained and undrained portions 45 years after the installation of a drainage ditch network, which has lowered water levels of 15–20 cm on average in the drained part of the site. Although shrub fine root production did not differ significantly between sites, water table drawdown increased belowground tree fine root production by 740% (3.8 ± 5.4 SD and 28.1 ± 24.1 g m^?2 y^?1 in undrained and drained sites, respectively) at the expense of herb root production, which declined 38% (27.62 ± 16.40 and 10.58 ± 15.7 g m^?2 y^?1 in undrained and drained sites, respectively) yielding no significant overall change in total fine root production. Drainage effects on aboveground biomass showed a similar pattern among plant types, as aboveground tree biomass increased dramatically with drainage (79 ± 135 and 2546 ± 1551 g m^?2 in drained and undrained sites, respectively). Although total shrub biomass was not significantly different between sites, shrubs allocated more biomass to stems than leaves in the drained site. Drainage also caused a significant shift in shrub species composition. Although trees dominated the aboveground biomass following water table drawdown, understorey vegetation, mainly shrubs, continued to dominate belowground fine root production, comprising 64% of total root production at the drained site. Aboveground biomass proved to be a good predictor of belowground production, suggesting that allometric relationships can be developed to estimate belowground production in these systems. Increase in tree root production can counteract decrease in herb fine root production following water table drawdown, emphasizing the importance of plant functional type responses to water table drawdown. Whether these changes will offset ecosystem C loss via increased plant C storage or stimulate soil organic matter decomposition via increased above- and belowground litter inputs requires further study.     

Phosphate Addition and Plant Species Alters Microbial Community Structure in Acidic Upland Grassland Soil

Agricultural improvement (addition of fertilizers, liming) of seminatural acidic grasslands across Ireland and the UK has resulted in significant shifts in floristic composition, soil chemistry, and microbial community structure. Although several factors have been proposed as responsible for driving shifts in microbial communities, the exact causes of such changes are not well defined. Phosphate was added to grassland microcosms to investigate the effect on fungal and bacterial communities. Plant species typical of unimproved grasslands (Agrostis capillaris, Festuca ovina) and agriculturally improved grasslands (Lolium perenne) were grown, and phosphate was added 25 days after seed germination, with harvesting after a further 50 days. Phosphate addition significantly increased root biomass (p < 0.001) and shoot biomass (p < 0.05), soil pH (by 0.1 U), and microbial activity (by 5.33 mg triphenylformazan [TPF] g⁻¹ soil; p < 0.001). A slight decrease (by 0.257 mg biomass-C g⁻¹ soil; p < 0.05) in microbial biomass after phosphate addition was found. The presence of plant species significantly decreased soil pH (p < 0.05; by up to 0.2 U) and increased microbial activity (by up to 6.02 mg TPF g⁻¹ soil) but had no significant effect on microbial biomass. Microbial communities were profiled using automated ribosomal intergenic spacer analysis. Multidimensional scaling plots and canonical correspondence analysis revealed that phosphate addition and its interactions with upland grassland plant species resulted in considerable changes in the fungal and bacterial communities of upland soil. The fungal community structure was significantly affected by both phosphate (R = 0.948) and plant species (R = 0.857), and the bacterial community structure was also significantly affected by phosphate (R = 0.758) and plant species (R = 0.753). Differences in microbial community structure following P addition were also revealed by similarity percentage analysis. These data suggest that phosphate application may be an important contributor to microbial community structural change during agricultural management of upland grasslands.     

基于物种多样性与生物量关系的草地群落稳定性对全球变暖的响应研究进展

全球变暖背景下物种多样性和生物量的空间变化格局及其对温度升高的响应是生态学研究的热点内容,是探究群落稳定性响应全球变暖的关键.物种分布是多个生态过程相互作用的产物,温度升高通过改变群落关键种的作用影响物种多样性,从而改变群落稳定性.关于草地群落稳定性对全球变暖的响应过程,国内外学者的相关研究存在很大差异.该文作者结合自...     

Effects of Drought and Warming on Biomass, Nutrient Allocation, and Oxidative Stress in Abies fabri in Eastern Tibetan Plateau

Abies fabri (Mast.) Craib is an endemic and dominant species in typical subalpine dark coniferous forests distributed in the eastern Tibetan Plateau. To assess how A. fabri may respond and adapt to future climate changes, we investigated the effects of drought and warming on the growth, resource allocation in biomass, membrane stability, and oxidative stress of the seedlings over two growing seasons. Drought (11.4 % average reduction in soil moisture) was created by excluding natural precipitation with a plastic roof and warming was performed by an infrared heater above the plots. Drought increased root length, the root-to-shoot ratio, N concentration, and N/P ratio in all organs, and decreased seedling height and C/N ratio in all organs. Moreover, warming (2 °C) decreased seedling height, root length, total biomass, and N concentration in stems but increased the C/N ratio. Furthermore, the combination of drought and warming decreased seedling height and biomass in all organs, which further increased the N concentration and N/P ratio in all organs. A significant decrease in the membrane stability index and an increase in malondialdehyde, superoxide radical (O₂ ^?), and hydrogen peroxide (H₂O₂) were exactly matched with a dramatic decrease of total biomass under the combination of drought and warming treatment. Together these results implied that drought alone and warming alone were unfavorable for the early growth of A. fabri, and drought plus warming will intensify the opposite effect of drought alone or warming alone. Moreover, N will be a limited nutrient under extant and future climate changes.     

Microclimatic Performance of a Free-Air Warming and CO2 Enrichment Experiment in Windy Wyoming,USA

In order to plan for global changing climate experiments are being conducted in many countries, but few have monitored the effects of the climate change treatments (warming, elevated CO₂) on the experimental plot microclimate. During three years of an eight year study with year-round feedback-controlled infra-red heater warming (1.5/3.0°C day/night) and growing season free-air CO₂ enrichment (600 ppm) in the mixed-grass prairie of Wyoming, USA, we monitored soil, leaf, canopy-air, above-canopy-air temperatures and relative humidity of control and treated experimental plots and evaluated ecologically important temperature differentials. Leaves were warmed somewhat less than the target settings (1.1 & 1.5°C day/night) but soil was warmed more creating an average that matched the target settings extremely well both during the day and night plus the summer and winter. The site typically has about 50% bare or litter covered soil, therefore soil heat transfer is more critical than in dense canopy ecosystems. The Wyoming site commonly has strong winds (5 ms^-1 average) and significant daily and seasonal temperature fluctuations (as much as 30°C daily) but the warming system was nearly always able to maintain the set temperatures regardless of abiotic variation. The within canopy-air was only slightly warmed and above canopy-air was not warmed by the system, therefore convective warming was minor. Elevated CO₂ had no direct effect nor interaction with the warming treatment on microclimate. Relative humidity within the plant canopy was only slightly reduced by warming. Soil water content was reduced by warming but increased by elevated CO₂. This study demonstrates the importance of monitoring the microclimate in manipulative field global change experiments so that critical physiological and ecological conclusions can be determined. Highly variable energy demand fluctuations showed that passive IR heater warming systems will not maintain desired warming for much of the time.     

灌丛化对黄土高原草地植物群落结构和地上生物量的影响

中国北方草地普遍出现灌丛化现象,灌丛化改变植物群落结构、植物多样性和生产力,直接影响着草地生态保护与可持续利用.该研究以黄土高原灌丛化草地为研究对象,通过植被调查,分析比较不同坡向的灌丛斑块与禾草斑块植物群落结构(物种组成、优势种及物种多样性)和地上生物量的差异.结果发现:(1)灌丛化草地不同坡向对物种多样性及地上生物...     

Effects of CO2Enrichment on Trees and Forests: Lessons to be Learned in View of Future Ecosystem Studies

Because of their prominent role in global bioproductivity andbecause of their complex structure and function, forests andtree species deserve particular attention in studies on thelikely impact of elevated atmospheric CO₂on terrestrial vegetation.Besides a synoptic review of some of the most prominent above-groundresponse processes, particular attention is given to below-groundresponses of trees to elevated atmospheric CO₂, while some feedbackprocesses and interactions with various biotic and abiotic factorsare also briefly summarized. At the leaf level there is littleevidence of the long-term loss of sensitivity to CO₂that wassuggested by earlier experiments with tree seedlings in pots.Future studies on photosynthesis measurements will probablynot alter our conclusions about acclimation, but should focusmore on respiration under elevated CO₂, which is still poorlyunderstood. At the tree level, the increase in growth observedin elevated CO₂results from an increase in both leaf area andleaf photosynthetic rate (per unit leaf area). Tree growth enhancementis generally larger at high rates of nutrient supply; when nutrientsupply rates do not meet growth rates, tree nutrient statusdeclines and nutrients become limiting. In many studies at thecanopy level, a shift in whole-tree carbon allocation patterntowards below-ground parts has been associated with increasedatmospheric CO₂concentrations. At the ecosystem level, a largeramount of carbon being allocated below-ground could show upby either (1) more root growth and turnover, (2) enhanced activityof root-associated microorganisms, (3) larger microbial biomasspools and enhanced microbial activity, or (4) increased lossesof soil carbon through soil respiration. Fine root productionis generally enhanced, but it is not clear whether this responsewould persist in a forest. As elevated CO₂stimulates biomassproduction, litterfall and rhizodeposition also increase. Thisincreased delivery of labile organic matter to the soil couldinfluence soil microbial communities and subsequent decompositionrates, nutrient availability and carbon storage in soil. Thereare, however, contradictory hypothesis about the direction inwhich nutrient availability will be affected. Knowledge of theresponse of these and other ecophysiological processes to elevatedCO₂is the key to understanding the functioning of the wholeforest ecosystem. Our current knowledge is sufficiently largewith regard to how the carbon uptake process and individualtree growth respond under atmospheric changes, but more emphasisshould be put in future experiments on the interactions betweenvarious processes, such as the carbon and nitrogen cycles, andon below-ground responses. Copyright 1999 Annals of Botany Company Global climatic changes, elevated CO₂, forests, trees, below-ground processes, mycorrhizae, roots, decomposition.     

Effects of Climate Change Drivers on Nitrous Oxide Fluxes in an Upland Temperate Grassland

Despite increasing interest in the patterns of trace gas emissions in terrestrial ecosystems, little is known about the impacts of climate change on nitrous oxide (N₂O) fluxes. The aim of this study was to determine the importance of the three main drivers of climate change (warming, summer drought, and elevated CO₂ concentrations) on N₂O fluxes from an extensively managed, upland grassland. Over a 2-year period, we monitored N₂O fluxes in an in situ ecosystem manipulation experiment simulating the climate predicted for the study area in 2080 (3.5°C temperature increase, 20% reduction in summer rainfall and atmospheric CO₂ levels of 600 ppm). N₂O fluxes showed significant seasonal and interannual variation irrespective of climate treatment, and were higher in summer and autumn compared with winter and spring. Overall, N₂O emissions showed a positive correlation with soil temperature and rainfall. Elevated temperature had a positive impact on mean annual N₂O fluxes but effects were only significant in 2007. Contrary to expectations, neither combined summer drought and warming nor the simultaneous application of elevated atmospheric CO₂ concentrations, summer drought and warming had any significant effect on annual N₂O fluxes. However, the maximum N₂O flux rates observed during the study occurred when elevated CO₂ was combined with warming and drought, suggesting the potential for important, short-term N₂O–N losses in enriched CO₂ environments. Taken together, our results suggest that the N₂O responses of temperate, extensively managed grasslands to future climate change scenarios may be primarily driven by temperature effects.     

Controls over Soil Nitrogen Pools in a Semiarid Grassland Under Elevated CO2 and Warming

Long-term responses of terrestrial ecosystems to the combined effects of warming and elevated CO₂ (eCO₂) will likely be regulated by N availability. The stock of soil N determines availability for organisms, but also influences loss to the atmosphere or groundwater. eCO₂ and warming can elicit changes in soil N via direct effects on microbial and plant activity, or indirectly, via soil moisture. Detangling the interplay of direct- and moisture-mediated impacts on soil N and the role of organisms in controlling soil N will improve predictions of ecosystem-level responses. We followed individual soil N pools over two growing seasons in a semiarid temperate grassland, at the Prairie Heating and CO₂ Enrichment experiment. We evaluated relationships of N pools with environmental factors and explored the role of plants by assessing plant biomass, plant N, and plant inputs to soil. We also assessed N forms in plots with and without vegetation to remove plant-mediated effects. Our study demonstrated that the effects of warming and eCO₂ are highly dependent on individual N form and on year. In this water-constrained grassland, eCO₂, warming and their combination appear to impact soil N pools through a complex combination of direct- and moisture-mediated effects. eCO₂ decreased NO₃ ^? but had neutral to positive effects on NH₄ ⁺ and dissolved organic N (DON), particularly in a wet year. Warming increased NO₃ ^? availability due to a combination of indirect drying and direct temperature-driven effects. Warming also increased DON only in vegetated plots, suggesting plant mediation. Our results suggest that impacts of combined eCO₂ and warming are not always equivalent for plant and soil pools; although warming can help offset the decrease in NO₃ ^? availability for plants under eCO₂, the NO₃ ^? pool in soil is mainly driven by the negative effects of eCO₂.