ࡱ> y{x[  bjbj "jjvl````RRRR$v, 4CCC      ! # ]C!"CCC ``s^    C`8 C j u:_, gg R <t 0 .$X$ ````Session 1: Crop Responses to Elevated CO2 Coordinator: Bruce Kimball Crop responses to free-air CO2 enrichment (FACE) experiments during the past 10 years are summarized. These FACE experiments were conducted on C3 grasses (wheat, perennial ryegrass, rice), a C4 grass (sorghum), a C3 legume (white clover), a C3 forb with tuber storage (potato), and C3 woody perennials (cotton, grape). Elevated CO2 increased photosynthesis, biomass, and yield substantially in C3 species, but little in C4. It decreased stomatal conductance in both C3 and C4 species and increased water-use efficiency (but did not necessarily reduce water use) in all crops. Growth stimulations were as large or larger under water-stress compared to well-watered conditions. At low soil N, stimulations of non-legumes were reduced, whereas elevated CO2 strongly stimulated growth of clover both at ample and low N conditions. Roots were generally stimulated more than shoots. Woody perennials had larger growth responses to elevated CO2 than herbs, but smaller reductions in stomatal conductance. Tissue N concentrations were lower while concentrations of carbohydrates and some other carbon-based compounds went up, with leaves being the organs affected most. Development was accelerated slightly in most species (due to warming brought about by stomatal closure?). Elevated CO2 affected some soil microbes greatly but not others (overall activity was stimulated). Detection of statistically significant changes in soil organic carbon in any one study was nearly impossible, but by combining results from several sites and years, it appeared that elevated CO2 increased soil carbon content. Comparisons of the FACE results with those from earlier chamber-based studies were consistent, which gives confidence that conclusions drawn from both types of experiment are accurate. Additional FACE and complementary experiments need to be conducted on other major crops (e.g., corn, barley, soybean), over a variety of climatic conditions. These experiments will be important for model development and testing. Combinations of experiments and models will provide improved assessments of effects of rising CO2 on crops. Such information is essential for policy makers concerned with environmental change. Session 2: The FACE Network Coordinators: Bob Nowak, Stan Smith An overview of the 18 non-agricultural FACE experiments will be presented that includes a summary of their location, design, and data collection effort. A potential value of having the network is the ability to test universal hypotheses concerning responses to elevated atmospheric CO2. An overview of several hypotheses, including those related to physiological processes, water and nitrogen availability, nutrient cycling, and biodiversity, will serve as examples of different network approaches. Because CO2 is often one of the most expensive on-going costs associated with FACE experiments, methods to conserve CO2 will be examined. Finally, the benefits of the current FACE experiments must be balanced against their necessarily artificial CO2-concentration step-change design. Session 3: Ecosystem and Physiological Adjustments to Elevated CO2 Coordinators: Evan DeLucia, Bert Drake, A.C. Finzi, Carla Gunderson, Yiqi Luo Increasing levels of atmospheric CO2 will stimulate photosynthesis and productivity in most ecosystems; the duration and magnitude of this stimulation is, however, an open question. Adjustments at the physiological, organism, and ecosystem levels have the potential to moderate the growth response to elevated CO2. We examined key feedbacks at different levels of organization that may influence the response to elevated CO2. The affect of acclimation to temperature on the carbon cycling also will be considered. At the physiological level, we will examine if the long-term responses of photosynthesis and respiration to elevated CO2 are predictable from the short-term responses. Most field studies show a sustained increase in photosynthesis even after years of growth in elevated CO2. There are, however, notable exceptions where the long-term response has been considerably smaller than expected. The patterns of physiological responses to CO2 with respect to taxonomy, development, leaf age, resource availability, sink activity, carbohydrate status, and environmental variation will be discussed. The growth response to elevated CO2 may be modulated at the organism-ecosystem level by changes in the relationship between the carbon and hydrologic cycles. Rising CO2 increases water use efficiency through a stimulation of photosynthesis and/or reduction of water loss.While it is difficult to identify the degree to which this stimulates NPP, it may be the most important factor increasing biomass accumulation in some ecosystems, particularly those limited by water. Another central question at the ecosystem level is whether the rate of N supply can meet the increased growth demands caused by elevated CO2. Exposure of a pine forest to an experimental increase in CO2 caused a stimulation of productivity and N uptake for the first five years of the experiment. However, there was no detectible increase in the rate of N mineralization from soil organic matter, or changes in nitrogen use efficiency or retranslocation efficiency of the forest. A decrease in the growth stimulation as this forest continues to exceed N demand is anticipated in the future. The potential interactions between elevated temperature and CO2 represents an important frontier in ecosystem research. The latest report by the Intergovernmental Panel on Climate Change predicts a 1.4-5.8 C average increase in the global surface temperature over the period 1990 to 2100. These estimates of warming are greater than earlier projections, partly because of the assumption that observed sensitivity of soil respiration to temperature under current climate conditions would hold in a warmer climate. A test of this assumption in a tall grass prairie ecosystem revealed that acclimation of soil respiration to warming might limit the positive feedback between the terrestrial carbon cycle and climate. Session 4: Enhancing Carbon Sequestration in Terrestrial Ecosystems Coordinators: Gary Jacobs, Blaine Metting The goal of this session is generate discussion on research that may lead to strategies for using terrestrial ecosystems as part of a technology portfolio to help slow the increase in atmospheric CO2 concentration. We will take stock of our understanding in plant and microbial systems, modeling, and carbon measurement technologies. From this, we will discuss research needs that can help improve estimates of the potential for increasing the storage of carbon through terrestrial ecosystem management and the technological and biological limitations to enhanced sequestration. A quick status report will be given on DOE projects for rapid and in situ/non-destructive soil carbon measurement. The session will use brief presentations summarizing recent research results and near- and long-term research needs. A brief discussion session at the end will focus on developing research plans to address the key future science needs and how to make our research relevant to decision and policy makers. The topics to be addressed are: (1) introduction: understanding, enhancing, and measuring carbon sequestration in terrestrial ecosystems; (2) challenges in plant science related to sequestration; (3) microbiological aspects of sequestration; (4) modeling soil carbon sequestration: status and future needs; (5) soil carbon measurement: summary of current DOE projects; (6) discussion and future science needs. Session 5: Testing Ecosystem Models with Independent Field Data Coordinators: Paul Hanson, Jeff Amthor Long-term field experiments and monitoring programs have provided databases appropriate for testing (evaluating) ecosystem models. Two recent multimodel testing exercises are highlighted. One exercise used 3 years of field data (ground and flux tower measurements) at the BOREAS Northern Old Black Spruce site (Manitoba, Canada) to evaluate nine ecosystem models with varying levels of physiological/ecological detail and time steps ranging from 30 minutes to one month. The other exercise, now in progress, is using 8 years of field data from the Oak Ridge (Tennessee) Throughfall Displacement Experiment site and nearby eddy covariance tower to test 13 models encompassing a range of ecological detail and model design criteria. In both exercises, model--model differences were apparent in most cases. Model--data agreement was good in some cases, but poor in many others. A key consideration (result) is that uncertainty in (or accuracy of) measurements is poorly known in many cases. This means that rigorous model tests are complicated at best, and often impossible. Near term (1-3 years) needs include improved methods of quantifying measurement uncertainty and broader application of methods of quantifying effects of model parameter uncertainty (or variation) on model output. Longer-term (5-10 years) needs are application of objective model testing techniques to multiple models using the large number of datasets being collected in field experiments. The most important information delivered in the next 1-3 years relevant to decision and policy makers will be improved understanding of capabilities (and limitations) of ecosystem models as tools for assessing effects of environmental changes on terrestrial ecosystem structure and functioning. Session 6: Elevated CO2 and Plant Respiration Coordinators: Steve Long, George Koch, Bert Drake Both instantaneous (or direct) and long-term (or indirect) effects of elevated CO2 on respiration have been reported. Two potential biochemical mechanisms have been proposed for the direct effect of CO2 on respiration: (1) direct inhibition of respiratory enzyme activity and (2) increased activity of dark CO2 fixation (e.g., via PEP carboxylase). The ability of these mechanisms to account for gas exchange observations is not supported by published studies, and recent experiments (including several measuring O2 rather than CO2 exchange) indicate that the direct effect on respiration is in part or whole an artifact of measuring CO2 exchange, specifically chamber leaks or incorrect use of CO2 analyzers, and most recently published studies on the subject have documented the lack of a direct effect. Moreover, a recent paper describes an alternative biological explanation of an apparent direct effect based on CO2 diffusion between the inside and outside of a gas exchange chamber via the intercellular air spaces of leaves. Just as is true for a leaky chamber gasket, a leaky leaf can give rise to an apparent direct effect when partial pressure of CO2 inside the chamber exceeds that in the air outside the chamber. We conclude that rising atmospheric CO2 will not significantly affect respiration through direct mechanisms. Mechanisms for an indirect effect of long-term elevated CO2 are more clear. (1) Tissue composition changes resulting from long-term elevated CO2 may influence growth or maintenance costs. (2) Changes in growth rate should cause corresponding changes in growth respiration rate. And (3) greater translocation from source leaves resulting from greater photosynthesis in elevated CO2 should increase respiration supporting translocation in those leaves. We examined Quercus geminata and Q. myrtifolia in Florida (OTCs ); Pinus taeda in N. Carolina (FACE) and; Populus tremuloides, Betula papyrifera, Acer saccharum in Wisconsin (FACE), grown at both elevated and ambient [CO2] for 5 years. No long-term decrease in respiration at elevated [CO2] was found in any case. After 5 years in elevated [CO2], P. tadea showed a significant 9% increase in respiration. In the scrub oak ecosystem, a significant 23% increase in respiration was evident in Q. geminata but not in the co-dominant Q. myrtifolia, after 5 years in elevated [CO2]. These studies not only confirm the growing view of no direct effect of elevated [CO2] on respiration, but also indicate that in the long-term there may be an increase in respiration per unit area and mass. Near-term needs: Further study of long-term effects of growth in elevated [CO2] on respiration. Mid-term needs: Validate use of respiratory O2 uptake as a surrogate for CO2 evolution. Determine whether CO2 treatments alter the respiratory quotient, and the conditions and bases for any alterations. Long-term needs: Understand the basis of increased respiration resulting from long-term elevated CO2 (i.e., relating respiration to the processes such as growth and transport that is supports). Session 7: Comparing Methods of Estimating NEP Coordinators: Peter Curtis, Paul Hanson, Steve Wofsy A comparison of eddy covariance estimates of annual NEP was made with estimates based on a combination of forest mensurational, ecophysiological, and other biometric methods at five AmeriFlux deciduous forest sites (across 10 of latitude and 18 of longitude, with similar stand ages, canopy heights, and stand basal areas). Annual aboveground NPP varied nearly two-fold among sites and was strongly correlated with average annual temperature and with estimated annual soil nitrogen mineralization (Nmin). Estimates of NEP ranged from a low of +0.3 Mg C ha-1 yr-1 in northern Michigan to a high of +3.5 Mg C ha-1 yr-1 in central Indiana, and were also correlated with Nmin. There was no systematic pattern among sites in over- versus under-estimation of the biometric compared to the tower-flux based estimates of NEP. Estimates of root and soil C dynamics were significant sources of uncertainty in biometric estimates of NEP and represent a prerequisite area of study needed for accurate estimates of forest net C storage. Interannual variation in amount of carbohydrate in storage (tree) pools has the potential to affect NEP by 100-200 g C m-2 y-1 in a given year (according to work at Oak Ridge, Tennessee), though this will likely be unimportant when averaged over multiple years. Application of ground based methods to estimate NEP for a Eucalyptus plantation in Hawaii that included real error estimates for all components (a critical step that is often ignored) revealed that average annual NEP over 6 years after pasture conversion was +10.3 Mg C/ha/yr for pastures fertilized at planting and +14.3 Mg C/ha/yr for pastures fertilized quarterly. NEP declined from +17.3 Mg C/ha/yr in year 1 to +7.6 Mg C/ha/yr in year 6. A new method of estimating soil CO2 flux at the catchment scale using stream water chemistry might be used to estimate ecosystem soil respiration independent of chamber and tower measurements, and may be an indicator of soil respiration at large spatial scales. These studies will provide decision and policy makers with a better understanding of uncertainty associated with tower-flux-based estimates of annual forest NEP. Session 8: Belowground Responses to Environmental Change: A Synthesis Coordinators: Don Zak, Julie Whitbeck, Hal Mooney The physiological activities of plant roots and soil organisms are important components of ecosystem responses to climatic change. In our session, we summarize how a changing environment will influence plant roots, mycorrhizal fungi, soil organisms, and soil C and N cycling. Key environmental change factors that we consider are rising atmospheric CO2 and O3, N deposition, and altered soil temperature and moisture (Fig. 1). An important conclusion of our synthesis is that plant growth and biomass allocation responses to the aforementioned factors cascade into the soil to directly modify the physiological activities of soil organisms and the ecosystem processes they mediate (i.e., soil C and N cycling). Experiments which directly manipulate climatic change factors have been critical for understanding the response of individual belowground components, and their results form the foundation for much of our current understanding of belowground responses. Our synthesis indicates that elevated CO2 and warmer soils generally increase allocation of photosynthate to roots and mycorrhizae, whereas high concentrations of ground-level ozone, N deposition, and drier soils lessen allocation to these structures. It is clear that changes in the activities of soil organisms follow these same trends, but we have an incomplete understanding of how changes in roots/mycorrhizae and the metabolism of soil organisms will interact to alter the supply of soil resources to plants. This remains an important ecosystem-level feedback for which we lack predictive power, and it is necessary to resolve this uncertainty to accurately predict how environmental change will alter the cycling and storage of C in terrestrial ecosystems. Session 9: Factors that Moderate the Response to Elevated CO2, and Natural Ecosystem Carbon Balance: Ozone and Nitrogen Coordinators: Dave Karnosky, Ram Oren, Herman Sievering Global atmospheric concentrations of CO2 and O3 have risen some 30 and 36% during the industrial period and they are rising concomitantly. Recent research suggests that elevated O3 offsets aboveground growth enhancement by elevated CO2 both for crops and forest trees. O3 also moderates belowground growth enhancement and microbial biomass production, alters plant responses to pests and affects processes cascading through the ecosystem. Results from these studies show that CO2 enrichment partially counteracts the deleterious effects of O3. However, maximum increases in agricultural crop and forest tree production in response to atmospheric CO2 enrichment in the future are likely to be jeopardized by concurrent increases in tropospheric O3 concentrations. Research at five FACE sites shows that: (1) In nutritionally rich sites (Aspen, WI; poplar, Italy), saplings grow faster under elevated CO2, diluting leaf N thus reducing carboxylation efficiency, so far with no effect on NPP. (2) In nutritionally moderate sites (Pine, NC; Sweetgum, TN) enhancement of C sequestration in woody biomass, primarily aboveground, lasted from 1 to 3 or 4 years, declining thereafter. Roughly at that time, foliar N concentration decreased, and C allocation to fine roots became more pronounced under elevated CO2. (3) In a nutritionally very poor site (prairie, MN), most species did not enhance growth aboveground with elevated CO2, but the enhancement was pronounced belowground. Aboveground biomass response to CO2 became noticeable for many species when N was added, a response also observed in whole-tree chambers with loblolly pine. Combining information from the pine FACE prototype (now running 8 years) with the full pine FACE experiment shows that more C and N accumulated in the forest floor and aboveground biomass under elevated CO2. Decreased N availability due to the initial surge in growth forced N uptake not to keep up with N demand, as also seen in sweetgum. Global C cycle assessments have suggested that atmospheric N deposition to forests may enhance photosynthesis and NEP. Carbon isotopic studies indicate a large C uptake in the 18(-53( N latitude zone where substantial anthropogenic N deposition occurs. However, direct experimental evidence, at the ecosystem or site scale, to support forest C uptake enhancement due to N deposition is limited. Data from the IBP Program indicate that both conifer and deciduous ecosystem aboveground growth are strongly dependent on foliar N requirement. This dependence on flux (N requirement) contrasts with a lack of dependence on foliar N content in the IBP study. Forest aboveground new growth, at the ecosystem scale, is not indicated to depend on the content of N in foliage as has been found at the leaf scale. Given the above, the atmospheric (including anthropogenic) deposition of N species to forest canopies may be a significant contributor to aboveground new growth. The foliar uptake of atmospheric N deposition together with N resorption and root uptake of N constitute the foliar N requirement for growth. At the Niwot AmeriFlux subalpine forest site, canopy uptake of atmospherically deposited N (mostly anthropogenic) has been observed to be 10-12% of aboveground N required for canopy growth. Studies at Whiteface Mountain indicate that anthropogenic N deposition to the forest canopy contributes about 10% of annual N requirement for growth. A recent study at the Howland forest is assessing eastern US forest C sequestration due to N additions to the forest canopy (by helicopter). Among the most important short-term (1-3 years) research needs are to (1) compare OTC and FACE results; (2) evaluate dose-responses of O3 and CO2 concurrently; (3) examine the mechanisms for CO2/O3 interactions with particular regard to antioxidants, and (4) conduct additional research on soil fertility effects on NPP and NEP under elevated CO2. Long-term (5-10 years) research needs include (1) compare CO2/O3 interactions of forest stands (LAI, WUE, biomass accumulation, etc.); (2) examine impacts of CO2/O3 on insect and diseases; (3) examine how ecosystem function (water and nutrient cycling, soil microorganisms, etc.) and determine how biodiversity is affected by these pollutants; and (4) studies of the long-term dynamics in the responses to elevated CO2, already seen in several studies, must be quantified. Information important to decision and policy makers that should be generated is on the interaction between site fertility, O3, and CO2 on productivity and C storage in pools of different longevity. Session 10: Carbon Cycle Modeling Coordinators: Mac Post, Scott Denning Session 11: The $-factor, Top-Down and Bottom-Up Coordinators: Dave Keeling, Rich Norby Models of the global fluxes of carbon between the atmosphere, ocean, and biosphere are used to understand and interpret the record of CO2 concentration in the atmosphere recorded at the Mauna Loa (Hawaii) Observatory. The $-factor, or biotic growth factor, is a first-order perturbation factor needed to describe the degree of plant growth stimulation in response to increasing CO2 concentration as it influences the flux between atmosphere and biosphere. Using current information on land use, the best fit to the atmospheric CO2 record occurs with $ set to 0.40 or 0.45 (with different assumptions in the ocean component of the model) compared to 0.35 in previous analyses. The response of net primary production in two ongoing FACE experiments yields somewhat higher, but generally comparable, values: 0.64 in the loblolly pine experiment at Duke Forest (North Carolina) and 0.53 in the deciduous sweetgum stand in Oak Ridge (Tennessee). A challenge for the near term will be to describe the experimentally derived $-factors in more robust terms that are more relevant to global carbon cycle models. More research is particularly needed on the permanence of the additional carbon taken up in elevated CO2. Within the next few years, an initial synthesis of FACE experiments and their implications for global models can be delivered. Session 12: Stable isotopes: contributions to the studies of ecosystem and carbon cycle research Coordinators: Jeff Chanton, Jim Ehleringer, Elise Pendall Stable isotopes at natural abundance levels provide a number of unique insights into the functioning of ecosystems and to improving our understanding of the carbon cycle. While most of the focus has been on carbon isotopes (13C/12C), analyses of oxygen (18O/16O) and nitrogen (15N/14N) are now adding new information about ecosystem functioning. At the regional and global scales, carbon and oxygen analyses of CO2 are critical for identifying the land/ocean partition of net CO2 uptake on annual and interannual bases. New studies are indicating that the isotopic signal emerging from ecosystems is not constant (as had been assumed), but fluctuates in response to water and other environmental factors. Integration of such biosphere-atmosphere studies provides a strong link between the ecosystem and global carbon cycle communities. At the ecosystem level, carbon and oxygen isotope analyses of CO2 are proving to be an integrating parameter for detecting ecosystem gas exchange responses to environmental changes. Stable isotope analyses also allow a partitioning of net ecosystem fluxes into their component fluxes, thereby providing a means for unraveling the independent responses of photosynthesis and respiration to environmental factors. Isotopic partitioning of these separate processes will allow more accurate modeling and future predictions of carbon sources and sinks. Within ecosystem scale elevated CO2 experiments (e.g., Free-Air CO2 Enrichment), the carbon isotope of the added CO2 is different from background values and provides a valuable tracer of new carbon input into the ecosystem. This tracer allows for quantification of different respiratory processes and for a quantitative analysis of carbon as it moves through trophic levels. Long-term observations of the isotopic composition of CO2 at the tropospheric and ecosystem scales are critical for partitioning the carbon fluxes at regional and global scales. When conducted over long time scales, this monitoring has significant implications for carbon-use and carbon-sequestration policies. Increased spatial coverage of these measurements, particularly in mid-continental areas, will reduce uncertainties in ecosystem processes in all regions of the globe. Session 13: Interannual Variation in Net Ecosystem Production (NEP) Coordinators: Dave Hollinger, Mike Goulden, Ken Davis Session 14: Spatial Representativeness of Flux Measurements/Models and Upscaling Carbon Fluxes to the Region Coordinators: HaPe Schmid, Walt Oechel, Jiquan Chen In order to upscale carbon fluxes to the region, a two-stage approach is required: the first stage requires quantification of the spatial representativeness of flux measurements, and the second is extrapolating these measurements to the region. To address these issues, footprint models that account for horizontal heterogeneity in the flow field, as well as sources/sinks that are distributed vertically need to be developed. The effects of topography on the spatial representativeness of flux measurements will also need considerable attention over the next decade. Scaling up from the flux footprint scale (or a single ecosystem type) to a mixed region can be achieved by combining (gridded) information about the biophysical characteristics of the surface with biosphere-atmosphere exchange models (one or three dimensional, dependent on the scale). One type of complication to the latter approach are highly fragmented landscapes because they lead to non-linear interactions across transitions. In these cases, flow models that can adequately account for advection are necessary for upscaling. At inhomogeneity length scales up to 103 m, higher order closure models or LES models have been shown to be effective tools. However, while LES resolves fluxes from seconds to hours (at best), ecological questions require flux estimates at annual or larger time scales. Future research into multiresolution and computational nesting approaches to adequately resolve such cross-scale transfer of information needs to be developed. An alternative to modeling of the scale integration is by use of spatially integrative flux measurements from either aircraft or dense networks of towers, but these face considerable problems of temporal or spatial coverage and cost. Over limited times and regions, such experimental scale integrations can be invaluable for model evaluation. At much larger spatial scales, inverse modeling methods, and the concept of virtual high towers, offer some promise to link troposheric CO2 concentration changes to surface fluxes, yet many unresolved issues remain. In the near future, methodologies for representativeness assessment and upscaling of surface fluxes need to be implemented across the network. The validation and implementation of these methodologies is instrumental for the experimental assessment of the regional or continental carbon budget. Session 15: Systematic Biases of Eddy Covariance Measurements of Carbon and/or Water Exchange Coordinator: Kell Wilson Bias errors are known to occur using the eddy covariance technique, but it is still uncertain how important bias errors are at all sites and if and how they should be corrected. Research has been focused in several directions: (1) diagnosing bias error, (2) understanding processes and conditions that lead to bias error, and (3) correcting for bias error. The diagnosis of bias error is being evaluated from studies of processing methods, spectral responses, independent measurements of the carbon budget, empirical relationships, energy balance closure, and most recently, from measurements that directly suggest additional non-zero terms in the equation for ecosystem exchange. Micrometeorological measurements have provided insight into the processes resulting in bias errors. Drainage flow, advection, mean persistent horizontal and vertical motions, and the pressure flux term have all been measured recently at Ameriflux sites. Atmospheric processes that may bias eddy covariance estimates have been identified at small and large scales. For example, drainage flows can occur very low to the ground and over fairly short distances. At larger scales, mean vertical and horizontal advection can occur because of land surface heterogeneity and induced thermal circulations. This has been shown to be true even over a flat surface with nearly homogeneous vegetation in the flux footprint, potentially redefining our definition of an ideal site. Correcting bias error has proceeded on three fronts: (1) the continued analysis of transfer functions (empirical and theoretical) and processing methods, (2) empirical relationships to gap fill, including the friction velocity threshold, and (3) direct estimates of additional terms in the flux equation. Future research should address the continued diagnosis of bias errors using various tools at more sites to examine how prevalent it is. Research should also indicate when empirical relationships are and are not appropriate and whether direct estimates of additional terms will provide the necessary quantitative precision.  Fig. 1 Atmospheric CO2 and O3, N deposition, and altered soil temperature and water regimes directly modify the amount of photosynthate allocated to roots and mycorrhizae. Changes in detritus production, N deposition, and modified soil temperature and water regimes directly alter the activities of soil organisms and the ecosystems processes they mediate (i.e., soil C & N cycling). 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