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Litter decay dynamics of paper birch (Betula papyrifera) were assessed at the Aspen free-air CO2 enrichment (FACE) facility in northern Wisconsin, USA. Leaf litter was decomposed for 12 months under factorial combinations of 360 vs. 560 μL CO2 L−1, crossed with 36 vs. 55 nL O3 L−1. To differentiate between substrate quality and environment effects, litterbags were placed in their Native Plots of origin or transplanted into the other treatments. CO2 enrichment, regardless of O3 concentration, produced poorer quality litter (high C/N, lignin/N and condensed tannins) than did ambient CO2 (low C/N, lignin/N and condensed tannins). Substrate quality differences were reflected in the mass loss rates (k-values), which were high for litter generated under ambient CO2 (0.887 year−1) and low for litter generated under elevated CO2 (0.674 year−1). The rate-retarding effects of CO2 enrichment were neither alleviated nor exacerbated by O3 exposure. Decay rates varied, however, depending on whether litter was placed back into its plot of origin or transplanted to Common Gardens. The results of this study are species specific, but they have important implications for understanding the processes regulating storage of fixed C and the release of CO2 from northern forest ecosystems. [ABSTRACT FROM AUTHOR]

Litter decay dynamics of paper birch (Betula papyrifera) were assessed at the Aspen free-air CO2 enrichment (FACE) facility in northern Wisconsin, USA. Leaf litter was decomposed for 12 months under factorial combinations of 360 vs. 560 μL CO2 L−1, crossed with 36 vs. 55 nL O3 L−1. To differentiate between substrate quality and environment effects, litterbags were placed in their Native Plots of origin or transplanted into the other treatments. CO2 enrichment, regardless of O3 concentration, produced poorer quality litter (high C/N, lignin/N and condensed tannins) than did ambient CO2 (low C/N, lignin/N and condensed tannins). Substrate quality differences were reflected in the mass loss rates (k-values), which were high for litter generated under ambient CO2 (0.887 year−1) and low for litter generated under elevated CO2 (0.674 year−1). The rate-retarding effects of CO2 enrichment were neither alleviated nor exacerbated by O3 exposure. Decay rates varied, however, depending on whether litter was placed back into its plot of origin or transplanted to Common Gardens. The results of this study are species specific, but they have important implications for understanding the processes regulating storage of fixed C and the release of CO2 from northern forest ecosystems. [ABSTRACT FROM AUTHOR]

The aim of this study was to examine the effects of elevated carbon dioxide [CO2] and ozone [O3] and their interaction on wood chemistry and anatomy of five clones of 3-year-old trembling aspen (Populus tremuloides Michx.). Wood chemistry was studied also on paper birch (Betula papyrifera Marsh.) and sugar maple (Acer saccharum Marsh.) seedling-origin saplings of the same age. Material for the study was collected from the Aspen Free-Air CO2 Enrichment (FACE) experiment in Rhinelander, WI, USA, where the saplings had been exposed to four treatments: control (C; ambient CO2, ambient O3), elevated CO2 (560ppm during daylight hours), elevated O3 (1.5 × ambient during daylight hours) and their combination (CO2 + O3) for three growing seasons (1998–2000). Wood chemistry responses to the elevated CO2 and O3 treatments differed between species. Aspen was most responsive, while maple was the least responsive of the three tree species. Aspen genotype affected the responses of wood chemistry and, to some extent, wood structure to the treatments. The lignin concentration increased under elevated O3 in four clones of aspen and in birch. However, elevated CO2 ameliorated the effect. In two aspen clones, nitrogen in wood samples decreased under combined exposure to CO2 and O2. Soluble sugar concentration in one aspen clone and starch concentration in two clones were increased by elevated CO2. In aspen wood, α-cellulose concentration changed under elevated CO2, decreasing under ambient 03 and slightly increasing under elevated O3. Hemicellulose concentration in birch was decreased by elevated CO2 and increased by elevated O3. In aspen, elevated O3 induced statistically significant reductions in distance from the pith to the bark and vessel lumen diameter, as well as increased wall thickness and wall percentage, and in one clone, decreased fibre lumen diameter. Our results show that juvenile wood properties of broadleaves, depending on species and genotype, were altered by atmospheric gas concentrations predicted for the year 2050 and that CO2 ameliorates some adverse effects of elevated O3 on wood chemistry. [ABSTRACT FROM AUTHOR]

This document is a response to concerns raised by Reershemius and Suhrhoff regarding a study conducted by Kantola et al. The authors of the response acknowledge that Reershemius and Suhrhoff did not find any faults with the derivation in the paper, which they state are factually true. They also mention that Reershemius and Suhrhoff arrived at a similar result in their own unpublished work. The response addresses specific implementation details raised by Reershemius and Suhrhoff and argues that these concerns lack merit. The authors appreciate the constructive engagement and transparency of data. [Extracted from the article]

Soil carbon sequestration is being considered as a potential pathway to mitigate climate change. Cropland soils could provide a sink for carbon that can be modified by farming practices; however, they can also act as a source of greenhouse gases (GHG), including not only nitrous oxide (N2O) and methane (CH4), but also the upstream carbon dioxide (CO2) emissions associated with agronomic management. These latter emissions are also sometimes termed "hidden" or "embedded" CO2. In this paper, we estimated the net GHG balance for Chinese cropping systems by considering the balance of soil carbon sequestration, N2O and CH4 emissions, and the upstream CO2 emissions of agronomic management from a life cycle perspective during 2000–2017. Results showed that although soil organic carbon (SOC) increased by 23.2 ± 8.6 Tg C per year, the soil N2O and CH4 emissions plus upstream CO2 emissions arising from agronomic management added 269.5 ± 21.1 Tg C‐eq per year to the atmosphere. These findings demonstrate that Chinese cropping systems are a net source of GHG emissions and that total GHG emissions are about 12 times larger than carbon uptake by soil sequestration. There were large variations between different cropping systems in the net GHG balance ranging from 328 to 7,567 kg C‐eq ha−1 year−1, but all systems act as a net GHG source to the atmosphere. The main sources of total GHG emissions are nitrogen fertilization (emissions during production and application), power use for irrigation, and soil N2O and CH4 emissions. Optimizing agronomic management practices, especially fertilization, irrigation, plastic mulching, and crop residues to reduce total GHG emissions from the whole chain is urgently required in order to develop a low‐carbon future for Chinese crop production. This is important to know what extent are greenhouse gas (GHG) emissions from intensive arable agriculture in China mitigated by sequestration of carbon in soil. We calculated the net GHG balance (kg C‐eq ha−1 year−1) between SOC stock change and emissions of N2O and CH4 and upstream CO2‐eq emissions associated with agronomic management. Despite the soil carbon sink found in Chinese cropland soils, it acted as a net GHG source because the emissions of total GHG are about one order of magnitude larger than the soil carbon sink under current farmers' practices. [ABSTRACT FROM AUTHOR]

We studied the three-way interaction of elevated CO2, nitrogen (N), and temperature ( T), and the two-way interaction of elevated CO2 and early-season defoliation on the secondary chemistry and resistance of Eurasian silver birch ( Betula pendula) and North American paper birch ( B. papyrifera) against the Eurasian hare ( Lepus timidus) and the North American eastern cottontail rabbit ( Sylvilagus floridanus), respectively. Elevated CO2 decreased the palatability of winter-dormant silver and paper birch stems to both hares and rabbits, respectively. But the effect on hares was only apparent at intermediate levels of N fertilization. Elevated T had no effect on palatability. The effects of elevated CO2, N, and T on levels of silver birch bark phenolics and terpenoids were dominated by two-way interactions between N and CO2, and N and T. Generally, however, N amendments elicited a parabolic response in carbon partitioning to most biosynthetic classes of silver birch phenolics (i.e. highest concentrations occurring at intermediate N). CO2 elevation was most enhancing at highest levels of N. On the other hand, T increases, more often than not, elicited reductions in phenolics, but especially so at the highest N level. In the case of B. papyrifera, elevated CO2 increased carbon partitioning to Folin-Denis stem and branch phenolics and condensed tannins. Early-season defoliation, on the other hand, had no effect on phenolics and tannins but lowered both N and energy levels of branches. Elevated CO2 substantially ameliorated the negative effects of severe defoliation on tree growth. These results support the hypothesis that continuing anthropogenic alterations of the atmosphere may trigger significant changes in plant phenotypic resistance to mammalian herbivores owing to an increasing net carbon balance between the highly vagile supply and demand capacities of plant carbon sources and sinks. [ABSTRACT FROM AUTHOR]

AbstractThis paper reviews two decades of effort by the scientific community in a search for predictive understanding of plant responses to elevated [CO2]. To evaluate the progress of research in leaf photosynthesis, plant respiration, root nutrient uptake, and carbon partitioning, we divided scientific activities into four phases: (I) initial assessments derived from our existing knowledge base to provide frameworks for experimental studies; (II) experimental tests of the initial assessments; (III) in cases where assessments were invalidated, synthesis of experimental results to stimulate alternative hypotheses and further experimentation; and (IV) formation of new knowledge. This paper suggests that photosynthetic research may have gone through all four phases, considering that (a) variable responses of photosynthesis to [CO2] are generally explainable, (b) extrapolation of leaf-level studies to the global scale has been examined, and (c) molecular studies are under way. Investigation of plant respiratory responses to [CO2] has reached the third phase: experimental results have been accumulated, and mechanistic approaches are being developed to examine alternative hypotheses in search for new concepts and/or new quantitative frameworks to understand respiratory responses to elevated [CO2]. The study of nutrient uptake kinetics is still in the second phase: experimental evidence has contradicted some of the initial assessments, and more experimental studies need to be designed before generalizations can be made. It is quite unfortunate that we have not made much progress in understanding mechanisms of carbon partitioning during the past two decades. This is due in part to the fact that some of the holistic theories, such as functional balance and optimality, have not evolved into testable hypotheses to guide experimental studies. This paper urges modelers to play an increasing role in plant–CO2 research by disassembling these existing theories into hypotheses and urges experimentalists to design experiments to examine these holistic concepts. [ABSTRACT FROM AUTHOR]

AbstractThis paper reviews two decades of effort by the scientific community in a search for predictive understanding of plant responses to elevated [CO2]. To evaluate the progress of research in leaf photosynthesis, plant respiration, root nutrient uptake, and carbon partitioning, we divided scientific activities into four phases: (I) initial assessments derived from our existing knowledge base to provide frameworks for experimental studies; (II) experimental tests of the initial assessments; (III) in cases where assessments were invalidated, synthesis of experimental results to stimulate alternative hypotheses and further experimentation; and (IV) formation of new knowledge. This paper suggests that photosynthetic research may have gone through all four phases, considering that (a) variable responses of photosynthesis to [CO2] are generally explainable, (b) extrapolation of leaf-level studies to the global scale has been examined, and (c) molecular studies are under way. Investigation of plant respiratory responses to [CO2] has reached the third phase: experimental results have been accumulated, and mechanistic approaches are being developed to examine alternative hypotheses in search for new concepts and/or new quantitative frameworks to understand respiratory responses to elevated [CO2]. The study of nutrient uptake kinetics is still in the second phase: experimental evidence has contradicted some of the initial assessments, and more experimental studies need to be designed before generalizations can be made. It is quite unfortunate that we have not made much progress in understanding mechanisms of carbon partitioning during the past two decades. This is due in part to the fact that some of the holistic theories, such as functional balance and optimality, have not evolved into testable hypotheses to guide experimental studies. This paper urges modelers to play an increasing role in plant–CO2 research by disassembling these existing theories into hypotheses and urges experimentalists to design experiments to examine these holistic concepts. [ABSTRACT FROM AUTHOR]

As human activity continues to increase CO2 and O3, broad expanses of north temperate forests will be simultaneously exposed to elevated concentrations of these trace gases. Although both CO2 and O3 are potent modifiers of plant growth, we do not understand the extent to which they alter competition for limiting soil nutrients, like nitrogen (N). We quantified the acquisition of soil N in two 8-year-old communities composed of trembling aspen genotypes ( n= 5) and trembling aspen–paper birch which were exposed to factorial combinations of CO2 (ambient and 560 μL L−1) and O3 (ambient = 30–40 vs. 50–60 nL L−1). Tracer amount of 15NH4+ were applied to soil to determine how these trace gases altered the competitive ability of genotypes and species to acquire soil N. One year after isotope addition, we assessed N acquisition by measuring the amount of 15N tracer contained in the plant canopy (i.e. recent N acquisition), as well as the total amount of canopy N (i.e. cumulative N acquisition). Exposure to elevated CO2 differentially altered recent and cumulative N acquisition among aspen genotypes, changing the rank order in which they obtained soil N. Elevated O3 also altered the rank order in which aspen genotypes obtained soil N by eliciting increases, decreases and no response among genotypes. If aspen genotypes respond similarly under field conditions, then rising concentrations of CO2 and O3 could alter the structure of aspen populations. In the aspen–birch community, elevated CO2 increased recent N (i.e. 15N) acquisition in birch (68%) to a greater extent than aspen (19%), suggesting that, over the course of this experiment, birch had gained a competitive advantage over aspen. The response of genotypes and species to rising CO2 and O3 concentrations, and how these responses are modified by competitive interactions, has the potential to change the future composition and productivity of northern temperate forests. [ABSTRACT FROM AUTHOR]

As human activity continues to increase CO2 and O3, broad expanses of north temperate forests will be simultaneously exposed to elevated concentrations of these trace gases. Although both CO2 and O3 are potent modifiers of plant growth, we do not understand the extent to which they alter competition for limiting soil nutrients, like nitrogen (N). We quantified the acquisition of soil N in two 8-year-old communities composed of trembling aspen genotypes ( n= 5) and trembling aspen–paper birch which were exposed to factorial combinations of CO2 (ambient and 560 μL L−1) and O3 (ambient = 30–40 vs. 50–60 nL L−1). Tracer amount of 15NH4+ were applied to soil to determine how these trace gases altered the competitive ability of genotypes and species to acquire soil N. One year after isotope addition, we assessed N acquisition by measuring the amount of 15N tracer contained in the plant canopy (i.e. recent N acquisition), as well as the total amount of canopy N (i.e. cumulative N acquisition). Exposure to elevated CO2 differentially altered recent and cumulative N acquisition among aspen genotypes, changing the rank order in which they obtained soil N. Elevated O3 also altered the rank order in which aspen genotypes obtained soil N by eliciting increases, decreases and no response among genotypes. If aspen genotypes respond similarly under field conditions, then rising concentrations of CO2 and O3 could alter the structure of aspen populations. In the aspen–birch community, elevated CO2 increased recent N (i.e. 15N) acquisition in birch (68%) to a greater extent than aspen (19%), suggesting that, over the course of this experiment, birch had gained a competitive advantage over aspen. The response of genotypes and species to rising CO2 and O3 concentrations, and how these responses are modified by competitive interactions, has the potential to change the future composition and productivity of northern temperate forests. [ABSTRACT FROM AUTHOR]

This paper examines long-term eddy covariance data from 18 European and 17 North American and Asian forest, wetland, tundra, grassland, and cropland sites under non-water-stressed conditions with an empirical rectangular hyperbolic light response model and a single layer two light-class carboxylase-based model. Relationships according to ecosystem functional type are demonstrated between empirical and physiological parameters, suggesting linkages between easily estimated parameters and those with greater potential for process interpretation. Relatively sparse documentation of leaf area index dynamics at flux tower sites is found to be a major difficulty in model inversion and flux interpretation. Therefore, a simplification of the physiological model is carried out for a subset of European network sites with extensive ancillary data. The results from these selected sites are used to derive a new parameter and means for comparing empirical and physiologically based methods across all sites, regardless of ancillary data. The results from the European analysis are then compared with results from the other Northern Hemisphere sites and similar relationships for the simplified process-based parameter were found to hold for European, North American, and Asian temperate and boreal climate zones. This parameter is useful for bridging between flux network observations and continental scale spatial simulations of vegetation/atmosphere carbon dioxide exchange. [ABSTRACT FROM AUTHOR]

Across many dryland regions, historically grass-dominated ecosystems have been encroached upon by woody-plant species. In this paper, we compare ecosystem water and carbon dioxide (CO2) fluxes over a grassland, a grassland–shrubland mosaic, and a fully developed woodland to evaluate potential consequences of woody-plant encroachment on important ecosystem processes. All three sites were located in the riparian corridor of a river in the southwest US. As such, plants in these ecosystems may have access to moisture at the capillary fringe of the near-surface water table. Using fluxes measured by eddy covariance in 2003 we found that ecosystem evapotranspiration (ET) and net ecosystem exchange of carbon dioxide (NEE) increased with increasing woody-plant dominance. Growing season ET totals were 407, 450, and 639 mm in the grassland, shrubland, and woodland, respectively, and in excess of precipitation by 227, 265, and 473 mm. This excess was derived from groundwater, especially during the extremely dry premonsoon period when this was the only source of moisture available to plants. Access to groundwater by the deep-rooted woody plants apparently decouples ecosystem ET from gross ecosystem production (GEP) with respect to precipitation. Compared with grasses, the woody plants were better able to use the stable groundwater source and had an increased net CO2 gain during the dry periods. This enhanced plant activity resulted in substantial accumulation of leaf litter on the soil surface that, during rainy periods, may lead to high microbial respiration rates that offset these photosynthetic fluxes. March–December (primary growing season) totals of NEE were −63, −212, and −233 g C m−2 in the grassland, shrubland, and woodland, respectively. Thus, there was a greater disparity between ecosystem water use and the strength of the CO2 sink as woody plants increased across the encroachment gradient. Despite a higher density of woody plants and a greater plant productivity in the woodland than in the shrubland, the woodland produced a larger respiration response to rainfall that largely offset its higher photosynthetic potential. These data suggest that the capacity for woody plants to exploit water resources in riparian areas results in enhanced carbon sequestration at the expense of increased groundwater use under current climate conditions, but the potential does not scale specifically as a function of woody-plant abundance. These results highlight the important roles of water sources and ecosystem structure on the control of water and carbon balances in dryland areas. [ABSTRACT FROM AUTHOR]

Net ecosystem production is the residual of two much larger fluxes: photosynthesis and respiration. While photosynthesis is a single process with a well-established theoretical underpinning, respiration integrates the variety of plant and microbial processes by which CO2 returns from ecosystems to the atmosphere. Limits to current capacity for predicting ecosystem respiration fluxes across biomes or years result from the mismatch between what is usually measured – bulk CO2 fluxes – and what process-based models can predict – fluxes of CO2 from plant (autotrophic) or microbial (heterotrophic) respiration. Papers in this Thematic Issue and in the recent literature, document advances in methods for separating respiration into autotrophic and heterotrophic components using three approaches: (1) continuous measurements of CO2 fluxes and assimilation of these data into process-based models; (2) application of isotope measurements, particularly radiocarbon; and (3) manipulation experiments. They highlight the role of allocation of C fixed by plants to respiration, storage, growth or transfer to other organisms as a control of seasonal and interannual variability in soil respiration and the oxidation state of C in the terrestrial biosphere. A second theme is the potential for comparing C isotope signatures in organic matter, CO2 evolved in incubations and microbial biomarkers to elucidate the pathways (respiration, recycling, or transformation) of C during decomposition. Together, these factors determine the continuum of timescales over which C is returned to the atmosphere by respiration and enable testing of theories of plant and microbial respiration that go beyond empirical models and allow predictions of future respiration responses to future change in climate, pollution and land use. [ABSTRACT FROM AUTHOR]

Dairy systems in Europe contribute to the emissions of the greenhouse gases (GHGs) nitrous oxide (N2O), methane (CH4) and carbon dioxide (CO2). In this paper, the effects of improved nitrogen (N) management on GHG emissions from Dutch dairy farms are determined. The GHG emissions are calculated using the panel on climate change (IPCC) methodology for the Netherlands, an updated and refined IPCC methodology, and a full accounting approach. The changes in dairy farming over the last 20 years, and the consequences for N management are described using detailed farm-level data, collected in 1985, 1997 and 2002. The selected years represent distinct stages in the implementation of N policies. The changes in N management have reduced the GHG emissions. A reduction of the N surplus per kilogram milk with 1 g N reduced the GHG emissions per kilogram milk with approximately 29 g CO2-equivalents. The reduction of the N surpluses was mainly brought about by reduced fertilizer use and reduced grazing time. The use of updated and refined emission factors resulted in higher CH4 emissions and lower N2O emissions. On average, the overall emission was 36% higher with the refined method. Full accounting, including all direct and indirect emissions of CH4, N2O and CO2, increased the emission with 36% compared with the refined IPCC methodology. We conclude that the N surplus at farm level is a useful indicator of GHG emissions. A full accounting system as presented in this study may effectively enable farmers to address the issue of emissions of GHGs in their operational management decisions. Both approaches serve their own specific objectives: full accounting at the farm level to explore mitigation options, and the IPCC methods to report changes in GHG emissions at the national level. [ABSTRACT FROM AUTHOR]

The effects of elevated carbon dioxide on plant–herbivore interactions have been summarized in a number of narrative reviews and metaanalyses, while accompanying elevation of temperature has not received sufficient attention. The goal of our study is to search, by means of metaanalysis, for a general pattern in responses of herbivores, and plant characteristics important for herbivores, to simultaneous experimental increase of carbon dioxide and temperature (ECET) in comparison with both ambient conditions and responses to elevated CO2 (EC) and temperature (ET) applied separately. Our database includes 42 papers describing studies of 31 plant species and seven herbivore species. Nitrogen concentration and C/N ratio in plants decreased under both EC and ECET treatments, whereas ET had no significant effect. Concentrations of nonstructural carbohydrates and phenolics increased in EC, decreased in ET and did not change in ECET treatments, whereas terpenes did not respond to EC but increased in both ET and ECET; leaf toughness increased in both EC and ECET. Responses of defensive secondary compounds to treatments differed between woody and green tissues as well as between gymnosperm and angiosperm plants. Insect herbivore performance was adversely affected by EC, favoured by ET, and not modified by ECET. Our analysis allowed to distinguish three types of relationships between CO2 and temperature elevation: (1) responses to EC do not depend on temperature (nitrogen, C/N, leaf toughness, phenolics in angiosperm leaves), (2) responses to EC are mitigated by ET (sugars and starch, terpenes in needles of gymnosperms, insect performance) and (3) effects emerge only under ECET (nitrogen in gymnosperms, and phenolics and terpenes in woody tissues). This result indicates that conclusions of CO2 elevation studies cannot be directly extrapolated to a more realistic climate change scenario. The predicted negative effects of CO2 elevation on herbivores are likely to be mitigated by temperature increase. [ABSTRACT FROM AUTHOR]

This paper presents results of 1 year (from March 25, 2003 to March 24, 2004, 366 days) of continuous measurements of net ecosystem CO2 exchange (NEE) above a steppe in Mongolia using the eddy covariance technique. The steppe, typical of central Mongolia, is dominated by C3 plants adapted to the continental climate. The following two questions are addressed: (1) how do NEE and its components: gross ecosystem production (GEP) and total ecosystem respiration ( Reco) vary seasonally? (2) how do NEE, GEP, and Reco respond to biotic and abiotic factors? The hourly minimal NEE and the hourly maximal Reco were −3.6 and 1.2 μmol m−2 s−1, respectively (negative values denoting net carbon uptake by the canopy from the atmosphere). Peak daily sums of NEE, GEP, and Reco were −2.3, 3.5, and 1.5 g C m−2 day−1, respectively. The annual sums of GEP, Reco, and NEE were 179, 138, and −41 g C m−2, respectively. The carbon removal by sheep was estimated to range between 10 and 82 g C m−2 yr−1 using four different approaches. Including these estimates in the overall carbon budget yielded net ecosystem productivity of −23 to +20 g C m−2 yr−1. Thus, within the remaining experimental uncertainty the carbon budget at this steppe site can be considered to be balanced. For the growing period (from April 23 to October 21, 2003), 26% and 53% of the variation in daily NEE and GEP, respectively, could be explained by the changes in leaf area index. Seasonality of GEP, Reco, and NEE was closely associated with precipitation, especially in the peak growing season when GEP and Reco were largest. Water stress was observed in late July to early August, which switched the steppe from a carbon sink to a carbon source. For the entire growing period, the light response curves of daytime NEE showed a rather low apparent quantum yield ( α=−0.0047 μmol CO2 μmol−1 photons of photosynthetically active radiation). However, the α values varied with air temperature ( Ta), vapor pressure deficit, and soil water content. [ABSTRACT FROM AUTHOR]

Modeling analyses suggest that an increase in growth rate of atmospheric CO2 concentrations during an anomalously warm year may be caused by a decrease in net ecosystem production (NEP) in response to increased heterotrophic respiration ( Rh). To test this hypothesis, 12 intact soil monoliths were excavated from a tallgrass prairie site near Purcell, Oklahoma, USA and divided among four large dynamic flux chambers (Ecologically Controlled Enclosed Lysimeter Laboratories (EcoCELLs)). During the first year, all four EcoCELLs were subjected to Oklahoma air temperatures. During the second year, air temperature in two EcoCELLs was increased by 4°C throughout the year to simulate anomalously warm conditions. This paper reports on the effect of warming on soil CO2 efflux, representing the sum of autotrophic respiration ( Ra) and Rh. During the pretreatment year, weekly average soil CO2 efflux was similar in all EcoCELLs. During the late spring, summer and early fall of the treatment year, however, soil CO2 efflux was significantly lower in the warmed EcoCELLs. In general, soil CO2 efflux was correlated with soil temperature and to a lesser extent with moisture. A combined temperature and moisture regression explained 64% of the observed variation in soil CO2 efflux. Soil CO2 efflux correlated well with a net primary production (NPP) weighted greenness index derived from digital photographs. Although separate relationships for control and warmed EcoCELLs showed better correlations, one single relationship explained close to 70% of the variation in soil CO2 efflux across treatments and years. A strong correlation between soil CO2 efflux and canopy development and the lack of initial response to warming indicate that soil CO2 efflux is dominated by Ra. This study showed that a decrease in soil CO2 efflux in response to a warm year was most likely dominated by a decrease in Ra instead of an increase in Rh. [ABSTRACT FROM AUTHOR]

This paper discusses the advantages and disadvantages of the different methods that separate net ecosystem exchange (NEE) into its major components, gross ecosystem carbon uptake (GEP) and ecosystem respiration ( Reco). In particular, we analyse the effect of the extrapolation of night-time values of ecosystem respiration into the daytime; this is usually done with a temperature response function that is derived from long-term data sets. For this analysis, we used 16 one-year-long data sets of carbon dioxide exchange measurements from European and US-American eddy covariance networks. These sites span from the boreal to Mediterranean climates, and include deciduous and evergreen forest, scrubland and crop ecosystems. We show that the temperature sensitivity of Reco, derived from long-term (annual) data sets, does not reflect the short-term temperature sensitivity that is effective when extrapolating from night- to daytime. Specifically, in summer active ecosystems the long-term temperature sensitivity exceeds the short-term sensitivity. Thus, in those ecosystems, the application of a long-term temperature sensitivity to the extrapolation of respiration from night to day leads to a systematic overestimation of ecosystem respiration from half-hourly to annual time-scales, which can reach >25% for an annual budget and which consequently affects estimates of GEP. Conversely, in summer passive (Mediterranean) ecosystems, the long-term temperature sensitivity is lower than the short-term temperature sensitivity resulting in underestimation of annual sums of respiration. We introduce a new generic algorithm that derives a short-term temperature sensitivity of Reco from eddy covariance data that applies this to the extrapolation from night- to daytime, and that further performs a filling of data gaps that exploits both, the covariance between fluxes and meteorological drivers and the temporal structure of the fluxes. While this algorithm should give less biased estimates of GEP and Reco, we discuss the remaining biases and recommend that eddy covariance measurements are still backed by ancillary flux measurements that can reduce the uncertainties inherent in the eddy covariance data. [ABSTRACT FROM AUTHOR]

Changes in atmospheric composition affect plant quality and herbivore performance. We used the Aspen Free Air CO2 Enrichment (FACE) facility to investigate the impacts of elevated carbon dioxide (CO2) and ozone (O3) on the performance of the aphid Cepegillettea betulaefoliae Granovsky feeding on paper birch ( Betula papyrifera Marsh.). In Year 1, we simultaneously measured individual performance and population growth rates, and in Year 2 we surveyed natural aphid, predator and parasitoid populations throughout the growing season. Aphid growth and development (relative growth rate (RGR), development time, adult weight, embryo number and the birth weight of newborn nymphs) were unaffected by CO2 and O3. Aphid fecundity decreased on trees grown at elevated CO2, O3 and CO2+O3. Neither nymphal performance nor adult size were reliable indicators of future fecundity at elevated CO2 and/or O3. Aphid populations protected from natural enemies were unaffected by elevated CO2, but increased significantly at elevated O3. Individual fecundity in elevated CO2 and O3 atmospheres did not predict population growth rates, probably because of changes in the strength of intraspecific competition or the ability of the aphids to induce nutrient sinks. Natural aphid, predator and parasitoids populations (Year 2) showed few significant responses to CO2 and O3, although CO2 and O3 did affect the timing of aphid and natural enemy peak abundance. Elevated CO2 and O3 affected aphid and natural enemy populations independently: no CO2× O3 interactions were observed. We conclude that: (1) aphid individual performance did not predict population responses to CO2 and O3 and (2) elevated CO2 and O3 atmospheres are unlikely to affect C. betulaefoliae populations in the presence of natural enemy communities. [ABSTRACT FROM AUTHOR]

Changes in atmospheric composition affect plant quality and herbivore performance. We used the Aspen Free Air CO2 Enrichment (FACE) facility to investigate the impacts of elevated carbon dioxide (CO2) and ozone (O3) on the performance of the aphid Cepegillettea betulaefoliae Granovsky feeding on paper birch ( Betula papyrifera Marsh.). In Year 1, we simultaneously measured individual performance and population growth rates, and in Year 2 we surveyed natural aphid, predator and parasitoid populations throughout the growing season. Aphid growth and development (relative growth rate (RGR), development time, adult weight, embryo number and the birth weight of newborn nymphs) were unaffected by CO2 and O3. Aphid fecundity decreased on trees grown at elevated CO2, O3 and CO2+O3. Neither nymphal performance nor adult size were reliable indicators of future fecundity at elevated CO2 and/or O3. Aphid populations protected from natural enemies were unaffected by elevated CO2, but increased significantly at elevated O3. Individual fecundity in elevated CO2 and O3 atmospheres did not predict population growth rates, probably because of changes in the strength of intraspecific competition or the ability of the aphids to induce nutrient sinks. Natural aphid, predator and parasitoids populations (Year 2) showed few significant responses to CO2 and O3, although CO2 and O3 did affect the timing of aphid and natural enemy peak abundance. Elevated CO2 and O3 affected aphid and natural enemy populations independently: no CO2× O3 interactions were observed. We conclude that: (1) aphid individual performance did not predict population responses to CO2 and O3 and (2) elevated CO2 and O3 atmospheres are unlikely to affect C. betulaefoliae populations in the presence of natural enemy communities. [ABSTRACT FROM AUTHOR]

Abstract This paper examines uncertainties in the interpretation of isotope signals when estimating fine root longevity, particularly in forests. The isotope signals are depleted δ13 C values from elevated CO2 experiments and enriched Δ14 C values from bomb 14 C in atmospheric CO2 . For the CO2 experiments, I explored the effects of six root mortality patterns (on–off, proportional, constant, normal, left skew, and right skew distributions), five levels of nonstructural carbohydrate (NSC) reserves, and increased root growth on root δ13 C values after CO2 fumigation. My analysis indicates that fitting a linear equation to δ13 C data provides unbiased estimates of longevity only if root mortality follows an on–off model, without dilution of isotope signals by pretreatment NSC reserves, and under a steady state between growth and death. If root mortality follows the other patterns, the linear extrapolation considerably overestimates root longevity. In contrast, fitting an exponential equation to δ13 C data underestimates longevity with all the mortality patterns except the proportional one. With either linear or exponential extrapolation, dilution of isotope signals by pretreatment NSC reserves could result in overestimation of root longevity by several-fold. Root longevity is underestimated if elevated CO2 stimulates fine root growth. For the bomb 14 C approach, I examined the effects of four mortality patterns (on–off, proportional, constant, and normal distribution) on root Δ14 C values. For a given Δ14 C value, the proportional pattern usually provides a shorter estimate of root longevity than the other patterns. Overall, we have to improve our understanding of root growth and mortality patterns and to measure NSC reserves in order to reduce uncertainties... [ABSTRACT FROM AUTHOR]

Abstract The free air carbon dioxide enrichment (FACE) and N deposition experiments on four ombrotrophic bogs in Finland, Sweden, the Netherlands and Switzerland, revealed that after three years of treatment: (1) elevated atmospheric CO2 concentration had no significant effect on the biomass growth of Sphagnum and vascular species; and (2) increased N deposition reduced Sphagnum growth, because it increased the cover of vascular plants and the tall moss Polytrichum strictum , while vascular plant biomass growth was not affected. This paper focuses on water chemistry, plant nutrient content, and litter decomposition rates. Potassium limitation, or low supply of K and P, may have prevented a significant increase of Sphagnum growth under elevated CO2 and N deposition. Vascular plant growth under elevated CO2 and N deposition was also limited by K, or by K in combination with P or N (N in CO2 experiment). Elevated CO2 and N deposition had no effect on decomposition rates of Sphagnum and vascular plant litter. Aside from a possible effect of N deposition on light competition between species, we expect that elevated atmospheric CO2 and N deposition concentrations will not affect Sphagnum and vascular plant growth in bogs of north-west Europe due to K-, or K in combination with N- or P-, limited growth. For the same reason we expect no effect of elevated CO2 and N deposition on litter decomposition. Net primary production of raised ombrotrophic bogs that are at or close to steady state, is regulated by input of nutrients through atmospheric deposition. Therefore, we hypothesize that the expected increase of plant growth under elevated CO2 and N deposition is diminished by current levels of K (and to some extent P and N) in atmospheric deposition. [ABSTRACT FROM AUTHOR]

Abstract This paper presents an empirical model of net ecosystem CO2 exchange (NEE) developed for a subarctic fen near Churchill, Manitoba. The model with observed data helps explain the interannual variability in growing season NEE. Five years of tower-flux data are used to test and examine the seasonal behaviour of the model simulations. Processes controlling the observed interannual variability of CO2 exchange at the fen are examined by exploring the sensitivity of the model to changes in air temperature, precipitation and leaf area index. Results indicate that the sensitivity of NEE to changing environmental controls is complex and varies interannually depending on the initial conditions of the wetland. Changes in air temperature and the timing of precipitation events have a strong influence on NEE, which is largely manifest in gross ecosystem photosynthesis (GEP). Climate change scenarios indicate that warmer air temperatures will increase carbon acquisition during wet years but may act to reduce wetland carbon storage in years that experience a large water deficit early in the growing season. Model simulations for this subarctic sedge fen indicate that carbon acquisition is greatest during wet and warm conditions. This suggests therefore that carbon accumulation was greatest at this subarctic fen during its early developmental stages when hydroclimatic conditions were relatively wet and warm at approximately 2500 years before present. [ABSTRACT FROM AUTHOR]

Summary Because biological and physical processes alter the stable isotopic composition of atmospheric CO2, variations in isotopic content can be used to investigate those processes. Isotopic flux measurements of 13CO2 above terrestrial ecosystems can potentially be used to separate net ecosystem CO2 exchange (NEE) into its component fluxes, net photosynthetic assimilation (FA) and ecosystem respiration (FR). In this paper theory is developed to partition measured NEE into FA and FR, using measurements of fluxes of CO2 and 13CO2, and isotopic composition of respired CO2 and forest air. The theory is then applied to fluxes measured (or estimated, for 13CO2) in a temperate deciduous forest in eastern Tennessee (Walker Branch Watershed). It appears that there is indeed enough additional information in 13CO2 fluxes to partition NEE into its photosynthetic and respiratory components. Diurnal patterns in FA and FR were obtained, which are consistent in magnitude and shape with patterns obtained from NEE measurements and an exponential regression between night-time NEE and temperature (a standard technique which provides alternate estimates of FR and FA). The light response curve for photosynthesis (FA vs. PAR) was weakly nonlinear, indicating potential for saturation at high light intensities. Assimilation-weighted discrimination against 13CO2 for this forest during July 1999 was 16.8–17.1‰, depending on canopy conductance. The greatest uncertainties in this approach lie in the evaluation of canopy conductance and its effect on whole-canopy photosynthetic discrimination, and thus the indirect methods used to estimate isotopic fluxes. Direct eddy covariance measurements of 13CO2 flux are needed to assess the validity of the assumptions used and provide defensible isotope-based estimates of the component fluxes of net ecosystem exchange. [ABSTRACT FROM AUTHOR]

Summary Because biological and physical processes alter the stable isotopic composition of atmospheric CO2, variations in isotopic content can be used to investigate those processes. Isotopic flux measurements of 13CO2 above terrestrial ecosystems can potentially be used to separate net ecosystem CO2 exchange (NEE) into its component fluxes, net photosynthetic assimilation (FA) and ecosystem respiration (FR). In this paper theory is developed to partition measured NEE into FA and FR, using measurements of fluxes of CO2 and 13CO2, and isotopic composition of respired CO2 and forest air. The theory is then applied to fluxes measured (or estimated, for 13CO2) in a temperate deciduous forest in eastern Tennessee (Walker Branch Watershed). It appears that there is indeed enough additional information in 13CO2 fluxes to partition NEE into its photosynthetic and respiratory components. Diurnal patterns in FA and FR were obtained, which are consistent in magnitude and shape with patterns obtained from NEE measurements and an exponential regression between night-time NEE and temperature (a standard technique which provides alternate estimates of FR and FA). The light response curve for photosynthesis (FA vs. PAR) was weakly nonlinear, indicating potential for saturation at high light intensities. Assimilation-weighted discrimination against 13CO2 for this forest during July 1999 was 16.8–17.1‰, depending on canopy conductance. The greatest uncertainties in this approach lie in the evaluation of canopy conductance and its effect on whole-canopy photosynthetic discrimination, and thus the indirect methods used to estimate isotopic fluxes. Direct eddy covariance measurements of 13CO2 flux are needed to assess the validity of the assumptions used and provide defensible isotope-based estimates of the component fluxes of net ecosystem exchange. [ABSTRACT FROM AUTHOR]

SummaryThe high-arctic environment is an environment where the consequences of global warming may be significant. In this paper we report on findings on carbon dioxide and water vapour fluxes above a sedge-dominated fen at Zackenberg (74°28′N, 20°34′ W) in The National Park of North and East Greenland. Eddy covariance measurements were initiated at the start of the growing season and terminated shortly before its end lasting 45 days. The net CO2 flux during daytime reaches a high of 10 μmol m–2s–1, and around the summer solstice, net CO2 assimilation occurred at midnight, resulting in net carbon gain during the night. The measured carbon dioxide fluxes compare well to estimates based on the photosynthesis model by Collatz et al. (1991). The total growing-season net ecosystem CO2 exchange was estimated to be 96 g C m–2 based on the carbon dioxide model and micrometeorological data. Finally, the combined CO2 assimilation and soil respiration models are used for examining the dependence of the carbon dioxide budget on temperature. The ecosystem is found to function optimally given the present temperature conditions whereas either an increase or a decrease in temperature would reduce the ecosystem CO2 accumulation. An increase in temperature by 5 °C would turn the ecosystem into a carbon dioxide source. [ABSTRACT FROM AUTHOR]

AbstractThe microclimate in facilities for studying effects of elevated CO2 on crops differs from ambient conditions. Open-top chambers (OTCs) increase temperature by 1–3 °C. If temperature and CO2 interact in their effect on crops, this would limit the value of OTC experiments. Furthermore, interaction of CO2 and temperature deserves study because increases in atmospheric CO2 concentration are expected to cause global warming. This paper describes two experiments in which a recently developed cooling system for OTCs was used to analyse the effects of temperature on photosynthesis, growth and yield of spring wheat (Triticum aestivum L., cv. Minaret). Two levels of CO2 were used (350 and 700 ppm), and two levels of temperature, with cooled OTCs being 1.6–2.4 °C colder than noncooled OTCs. Photosynthetic rates were increased by elevated CO2, but no effect of temperature was found. Cross-switching CO2 concentrations as well as determination of A–Ci curves showed that plant photosynthetic capacity after anthesis acclimated to elevated CO2. The acclimation may be related to the effects of CO2 on tissue composition: elevated CO2 decreased leaf nitrogen concentrations and increased sugar content. Calculations of the seasonal mean crop light-use efficiency (LUE) were consistent with the photosynthesis data in that CO2 increased LUE by 20% on average whereas temperature had no effect. Both elevating CO2 and cooling increased grain yield, by an average of 11% and 23%, respectively. CO2 and temperature stimulated yield via different mechanisms: CO2 increased photosynthetic rate, but decreased crop light interception capacity (LAI), whereas cooling increased grain yield by increasing LAI and extending the growing season with 10 days. The effects of CO2 and temperature were not additive: the CO2 effect was about doubled in the noncooled open-top chambers. In most cases, effects on yield were mediated through increased grain density rather than increased individual grain weights. The higher growth response to elevated CO2 in noncooled vs. cooled OTCs shows that a cooling system may remove a bias towards overestimating crop growth response to CO2 in open-top chambers. [ABSTRACT FROM AUTHOR]

The use of fossil fuel is predicted to cause an increase of the atmospheric CO2 concentration, which will affect the global pattern of temperature and precipitation. It is therefore essential to incorporate effects of temperature and water supply on carbon partitioning of plants to predict effects of elevated [CO2] on growth and yield of Triticum aestivum. Although earlier papers have emphasized that elevated [CO2] favours investment of biomass in roots relative to that in leaves, it has now become clear that these are indirect effects, due to the more rapid depletion of nutrients in the root environment as a consequence of enhanced growth. Broadly generalized, the effect of temperature on biomass allocation in the vegetative stage is that the relative investment of biomass in roots is lowest at a certain optimum temperature and increases at both higher and lower temperatures. This is found not only when the temperature of the entire plant is varied, but also when only root temperature is changed whilst shoot temperature is kept constant. Effects of temperature on the allocation pattern can be explained largely by the effect of root temperature on the roots' capacity to transport water. Effects of a shortage in water supply on carbon partitioning are unambiguous: roots receive relatively more carbon. The pattern of biomass allocation in the vegetative stage and variation in water-use efficiency are prime factors determining a plant's potential for early growth and yield in different environments. In a comparison of a range of T. aestivum cultivars, a high wateruse efficiency at the plant level correlates positively with a large investment in both leaf and root biomass, a low stomatal conductance and a large investment in photosynthetic capacity. We also present evidence that a lower investment of biomass in roots is not only associated with lower respiratory costs for root growth, but also with lower specific costs for ion uptake. We suggest the combination of a number of traits in future wheat cultivars, i.e. a high investment of biomass in leaves, which have a low stomatal conductance and a high photosynthetic capacity, and a low investment of biomass in roots, which have low respiratory costs. Such cultivars are considered highly appropriate in a future world, especially in the dryer regions. Although variation for the desired traits already exists among wheat cultivars, it is much larger among wild Aegilops species, which can readily be crossed with T. aestivum. Such wild relatives may be exploited to develop new wheat cultivars well-adapted to changed climatic conditions. [ABSTRACT FROM AUTHOR]

Air temperature and the atmospheric concentrations of carbon dioxide are expected to rise. These two factor have a great potential to affect development, growth and yield of crops, including wheat. Rising air temperature may affect wheat development more than rising atmospheric CO2 as there is not yet evidence that elevated CO2 concentrations can directly induce changes in wheat development. In winter wheat, temperature has a complex effect on development due to its strong interaction with vernalization and photoperiod. In this paper, potential effects of rising temperature on the development of winter wheat from sowing to heading are considered in the light of this complex controlling mechanism. Data from a large series of field trials made in Romania is analysed at first and, subsequently, the IATA-Wheat Phenology model is used to calculate the impact of air warming on wheat development under different climate change scenarios. Data from the field trials showed very clearly the occurrence of a complex temperature/photoperiod/vernalization interaction for field sown crops and demostrated that the photoperiodic and vernalization responses have a key role in controlling the duration of the emergence-heading period. Temperature plays, instead, a central role in controlling seed germination and crop emergence as well as leaf inititiation and leaf appearance rate. The results of model analysis showed very well that the impact of an even or uneven distribution of warming effects may be very different. In the first case, the model predicted that the duration of the vegetative period was at least partly reduced in some years. In the second case, the model suggested that if warming will be more pronounced in winter than in spring, as predicted for some areas of the world by General Circulation Models, we may expect an increase in the duration of the vegetative phase of growth. On the contrary, in case of a spring warming but unchanged winter temperatures, we may expect a substantial decrease in the duration of the vegetative period. [ABSTRACT FROM AUTHOR]

Free Air CO2 Enrichment (FACE) systems are used to fumigate unconfined field plots with CO2. As these installationscan treat a sufficiently large area without interfering with naturalclimatic conditions, they are considered important tools for global change research worldwide. However, there is general consensus that elevated capital costs of existing FACE systems as well as high running costs may prevent their application at the required level of scale.A new and small FACE system that was designed to reduce both capitalcosts and CO2 use, is described in this paper. Due to itsintermediate size (8 m diameter) between the smaller Mini-FACE systems that were developed in Italy and the larger systems designed by the Brookhaven National Laboratory in the USA, it was named Mid-FACE. The Mid-FACE was at first developed as a prototype and then used to enrich field grown potato crops in a CO2 concentration gradient experimental design. Technical details of a Mid-FACE prototype and of the operational set-up are presented in this paper together withperformance data in terms of temporal and spatial control of CO2 concentrations within the experimental area. [ABSTRACT FROM AUTHOR]