Beyond carbon flux partitioning: Carbon allocation and nonstructural carbon dynamics inferred from continuous fluxes.

Carbon (C) allocation and nonstructural carbon (NSC) dynamics play essential roles in plant growth and survival under stress and disturbance. However, quantitative understanding of these processes remains limited. Here we propose a framework where we connect commonly measured carbon cycle components (eddy covariance fluxes of canopy CO2 exchange, soil CO2 efflux, and allometry-based biomass and net primary production) by a simple mass balance model to derive ecosystem-level NSC dynamics (NSCi ), C translocation (dCi ), and the biomass production efficiency (BPEi ) in above- and belowground plant (i = agp and bgp) compartments. We applied this framework to two long-term monitored loblolly pine (Pinus taeda) plantations of different ages in North Carolina and characterized the variations of NSC and allocation in years under normal and drought conditions. The results indicated that the young stand did not have net NSC flux at the annual scale, whereas the mature stand stored a near-constant proportion of new assimilates as NSC every year under normal conditions, which was comparable in magnitude to new structural growth. Roots consumed NSC in drought and stored a significant amount of NSC post drought. The above- and belowground dCi and BPEi varied more from year to year in the young stand and approached a relatively stable pattern in the mature stand. The belowground BPEbgp differed the most between the young and mature stands and was most responsive to drought. With the internal C dynamics quantified, this framework may also improve biomass production estimation, which reveals the variations resulting from droughts. Overall, these quantified ecosystem-scale dynamics were consistent with existing evidence from tree-based manipulative experiments and measurements and demonstrated that combining the continuous fluxes as proposed here can provide additional information about plant internal C dynamics. Given that it is based on broadly available flux data, the proposed framework is promising to improve the allocation algorithms in ecosystem C cycle models and offers new insights into observed variability in soil-plant-climate interactions.

Spatial variability in tree-ring carbon isotope discrimination in response to local drought across the entire loblolly pine natural range.

Considering the temporal responses of carbon isotope discrimination (Δ13C) to local water availability in the spatial analysis of Δ13C is essential for evaluating the contribution of environmental and genetic facets of plant Δ13C. Using tree-ring Δ13C from years with contrasting water availability at 76 locations across the natural range of loblolly pine, we decomposed site-level Δ13C signals to maximum Δ13C in well-watered conditions (Δ13Cmax) and isotopic drought sensitivity (m) as a change in Δ13C per unit change of Palmer's Drought Severity Index (PDSI). Site water status, especially the tree lifetime average PDSI, was the primary factor affecting Δ13Cmax. The strong spatial correlation exhibited by m was related to both genetic and environmental factors. The long-term average water availability during the period relevant to trees as indicated by lifetime average PDSI correlated with Δ13Cmax, suggesting acclimation in tree gas-exchange traits, independent of incident water availability. The positive correlation between lifetime average PDSI and m indicated that loblolly pines were more sensitive to drought at mesic than xeric sites. The m was found to relate to a plant's stomatal control and may be employed as a genetic indicator of efficient water use strategies. Partitioning Δ13C to Δ13Cmax and m provided a new angle for understanding sources of variation in plant Δ13C, with several fundamental and applied implications.

Aquaporins, and not changes in root structure, provide new insights into physiological responses to drought, flooding, and salinity.

The influence of aquaporin (AQP) activity on plant water movement remains unclear, especially in plants subject to unfavorable conditions. We applied a multitiered approach at a range of plant scales to (i) characterize the resistances controlling water transport under drought, flooding, and flooding plus salinity conditions; (ii) quantify the respective effects of AQP activity and xylem structure on root (Kroot), stem (Kstem), and leaf (Kleaf) conductances; and (iii) evaluate the impact of AQP-regulated transport capacity on gas exchange. We found that drought, flooding, and flooding plus salinity reduced Kroot and root AQP activity in Pinus taeda, whereas Kroot of the flood-tolerant Taxodium distichum did not decline under flooding. The extent of the AQP control of transport efficiency varied among organs and species, ranging from 35-55% in Kroot to 10-30% in Kstem and Kleaf. In response to treatments, AQP-mediated inhibition of Kroot rather than changes in xylem acclimation controlled the fluctuations in Kroot. The reduction in stomatal conductance and its sensitivity to vapor pressure deficit were direct responses to decreased whole-plant conductance triggered by lower Kroot and larger resistance belowground. Our results provide new mechanistic and functional insights on plant hydraulics that are essential to quantifying the influences of future stress on ecosystem function.

An improved method for quantifying total fine root decomposition in plantation forests combining measurements of soil coring and minirhizotrons with a mass balance model.

Accurate measurement of total fine root decomposition (the amount of dead fine roots decomposed per unit soil volume) is essential for constructing a soil carbon budget. However, the ingrowth/soil core-based models are dependent on the assumptions that fine roots in litterbags/intact cores have the same relative decomposition rate as those in intact soils and that fine root growth and death rates remain constant over time, while minirhizotrons cannot quantify the total fine root decomposition. To improve the accuracy of estimates for total fine root decomposition, we propose a new method (balanced hybrid) with two models that integrate measurements of soil coring and minirhizotrons into a mass balance model. Model input parameters were fine root biomass, necromass and turnover rate for Model 1, and fine root biomass, necromass and death rate for Model 2. We tested the balanced hybrid method in a loblolly pine plantation forest in coastal North Carolina, USA. The total decomposition rate of absorptive fine roots (ARs) (a combination of first- and second-order fine roots) using Models 1 and 2 was 107 ± 13 g m-2 year-1 and 129 ± 12 g m-2 year-1, respectively. Monthly total AR decomposition was highest from August to November, which corresponded with the highest monthly total ARs mortality. The ARs imaged by minirhizotrons well represent those growing in intact soils, evident by a significant and positive relationship between the standing biomass and the standing length. The total decomposition estimate in both models was sensitive to changes in fine root biomass, turnover rate and death rate but not to change in necromass. Compared with Model 2, Model 1 can avoid the technical difficulty of deciding dead time of individual fine roots but requires greater time and effort to accurately measure fine root biomass dynamics. The balanced hybrid method is an improved technique for measuring total fine root decomposition in plantation forests in which the estimates are based on empirical data from soil coring and minirhizotrons, moving beyond assumptions of traditional approaches.

Using δ13C and δ18O to analyze loblolly pine (Pinus taeda L.) response to experimental drought and fertilization.

Drought frequency and intensity are projected to increase throughout the southeastern USA, the natural range of loblolly pine (Pinus taeda L.), and are expected to have major ecological and economic implications. We analyzed the carbon and oxygen isotopic compositions in tree ring cellulose of loblolly pine in a factorial drought (~30% throughfall reduction) and fertilization experiment, supplemented with trunk sap flow, allometry and microclimate data. We then simulated leaf temperature and applied a multi-dimensional sensitivity analysis to interpret the changes in the oxygen isotope data. This analysis found that the observed changes in tree ring cellulose could only be accounted for by inferring a change in the isotopic composition of the source water, indicating that the drought treatment increased the uptake of stored moisture from earlier precipitation events. The drought treatment also increased intrinsic water-use efficiency, but had no effect on growth, indicating that photosynthesis remained relatively unaffected despite 19% decrease in canopy conductance. In contrast, fertilization increased growth, but had no effect on the isotopic composition of tree ring cellulose, indicating that the fertilizer gains in biomass were attributable to greater leaf area and not to changes in leaf-level gas exchange. The multi-dimensional sensitivity analysis explored model behavior under different scenarios, highlighting the importance of explicit consideration of leaf temperature in the oxygen isotope discrimination (Δ18Oc) simulation and is expected to expand the inference space of the Δ18Oc models for plant ecophysiological studies.

An extractive removal step optimized for a high-throughput α-cellulose extraction method for δ13C and δ18O stable isotope ratio analysis in conifer tree rings.

Stable isotope ratios (δ13C and δ18O) of tree-ring α-cellulose are important tools in paleoclimatology, ecology, plant physiology and genetics. The Multiple Sample Isolation System for Solids (MSISS) was a major advance in the tree-ring α-cellulose extraction methods, offering greater throughput and reduced labor input compared to traditional alternatives. However, the usability of the method for resinous conifer species may be limited by the need to remove extractives from some conifer species in a separate pretreatment step. Here we test the necessity of pretreatment for α-cellulose extraction in loblolly pine (Pinus taeda L.), and the efficiency of a modified acetone-based ambient-temperature step for the removal of extractives (i) in loblolly pine from five geographic locations representing its natural range in the southeastern USA, and (ii) on five other common coniferous species (black spruce (Picea mariana Mill.), Fraser fir (Abies fraseri (Pursh) Poir.), Douglas fir (Pseudotsuga menziesii (Mirb.) Franco), Norway spruce (Picea abies (L.) Karst) and ponderosa pine (Pinus ponderosa D.)) with contrasting extractive profiles. The differences of δ13C values between the new and traditional pretreatment methods were within the precision of the isotope ratio mass spectrometry method used (±0.2‰), and the differences between δ18O values were not statistically significant. Although some unanticipated results were observed in Fraser fir, the new ambient-temperature technique was deemed as effective as the more labor-consuming and toxic traditional pretreatment protocol. The proposed technique requires a separate acetone-inert multiport system similar to MSISS, and the execution of both pretreatment and main extraction steps allows for simultaneous treatment of up to several hundred microsamples from resinous softwood, while the need of additional labor input remains minimal.

Hemlock woolly adelgid (Adelges tsugae) infestation affects water and carbon relations of eastern hemlock (Tsuga canadensis) and Carolina hemlock (Tsuga caroliniana).

Hemlock woolly adelgid (HWA) is an exotic insect pest causing severe decimation of native hemlock trees. Extensive research has been conducted on the ecological impacts of HWA, but the exact physiological mechanisms that cause mortality are not known. Water relations, anatomy and gas exchange measurements were assessed on healthy and infested eastern (Tsuga canadensis) and Carolina (Tsuga caroliniana) hemlock trees. These data were then used in a mechanistic model to test whether the physiological responses to HWA infestation were sufficiently significant to induce changes in whole-plant water use and carbon uptake. The results indicated coordinated responses of functional traits governing water relations in infested relative to healthy trees. In response to HWA, leaf water potential, carbon isotope ratios, plant hydraulic properties and stomatal conductance were affected, inducing a reduction in tree water use by > 40% and gross primary productivity by 25%. Anatomical changes also appeared, including the activation of traumatic cells. HWA infestation had a direct effect on plant water relations. Despite some leaf compensatory mechanisms, such as an increase in leaf hydraulic conductance and nitrogen content, tree water use and carbon assimilation were diminished significantly in infested trees, which could contribute to tree mortality.

Interactive effects of nocturnal transpiration and climate change on the root hydraulic redistribution and carbon and water budgets of southern United States pine plantations.

Deep root water uptake and hydraulic redistribution (HR) have been shown to play a major role in forest ecosystems during drought, but little is known about the impact of climate change, fertilization and soil characteristics on HR and its consequences on water and carbon fluxes. Using data from three mid-rotation loblolly pine plantations, and simulations with the process-based model MuSICA, this study indicated that HR can mitigate the effects of soil drying and had important implications for carbon uptake potential and net ecosystem exchange (NEE), especially when N fertilization is considered. At the coastal site (C), characterized by deep organic soil, HR increased dry season tree transpiration (T) by up to 40%, and such an increase affected NEE through major changes in gross primary productivity (GPP). Deep-rooted trees did not necessarily translate into a large volume of HR unless soil texture allowed large water potential gradients to occur, as was the case at the sandy site (S). At the Piedmont site (P) characterized by a shallow clay-loam soil, HR was low but not negligible, representing up to 10% of T. In the absence of HR, it was predicted that at the C, S and P sites, annual GPP would have been diminished by 19, 7 and 9%, respectively. Under future climate conditions HR was predicted to be reduced by up to 25% at the C site, reducing the resilience of trees to precipitation deficits. The effect of HR on T and GPP was predicted to diminish under future conditions by 12 and 6% at the C and P sites, respectively. Under future conditions, T was predicted to stay the same at the P site, but to be marginally reduced at the C site and slightly increased at the S site. Future conditions and N fertilization would decrease T by 25% at the C site, by 15% at the P site and by 8% at the S site. At the C and S sites, GPP was estimated to increase by 18% and by >70% under future conditions, respectively, with little effect of N fertilization. At the P site, future conditions would stimulate GPP by only 12%, but future conditions plus N fertilization would increase GPP by 24%. As a consequence, in all sites, water use efficiency was predicted to improve dramatically with future conditions. Modeling the effect of reduced annual precipitation indicated that limited water availability would decrease all carbon fluxes, including NEE and respiration. Our simulations highlight the interactive effects of nutrients and elevated CO(2), and showed that the effect of N fertilization would be greater under future climate conditions.

The role of harvest residue in rotation cycle carbon balance in loblolly pine plantations. Respiration partitioning approach.

Timber harvests remove a significant portion of ecosystem carbon. While some of the wood products moved off-site may last past the harvest cycle of the particular forest crop, the effect of the episodic disturbances on long-term on-site carbon sequestration is unclear. The current study presents a 25 year carbon budget estimate for a typical commercial loblolly pine plantation in North Carolina, USA, spanning the entire rotation cycle. We use a chronosequence approach, based on 5 years of data from two adjacent loblolly pine plantations. We found that while the ecosystem is very productive (GEP up to 2900 g m-2  yr-1 , NEE at maturity about 900 g C m-2  yr-1 ), the production of detritus does not offset the loss of soil C through heterotrophic respiration (RH ) on an annual basis. The input of dead roots at harvest may offset the losses, but there remain significant uncertainties about both the size and decomposition dynamics of this pool. The pulse of detritus produced at harvest resulted in a more than 60% increase in RH . Contrary to expectations, the peak of RH in relation to soil respiration (SR) did not occur immediately after the harvest disturbance, but in years 3 and 4, suggesting that a pool of roots may have remained alive for the first few years. On the other hand, the pulse of aboveground RH from coarse woody debris lasted only 2 years. The postharvest increase in RH was offset by a decrease in autotrophic respiration such that the total ecosystem respiration changed little. The observed flux rates show that even though the soil C pool may not necessarily decrease in the long-term, old soil C is definitely an active component in the site C cycle, contributing about 25-30% of the RH over the rotation cycle.

Genetic effects on total phenolics, condensed tannins and non-structural carbohydrates in loblolly pine (Pinus taeda L.) needles.

Carbon allocation to soluble phenolics (total phenolics, proanthocyanidins (PA)) and total non-structural carbohydrates (TNC; starch and soluble sugars) in needles of widely planted, highly productive loblolly pine (Pinus taeda L.) genotypes could impact stand resistance to herbivory, and biogeochemical cycling in the southeastern USA. However, genetic and growth-related effects on loblolly pine needle chemistry are not well characterized. Therefore, we investigated genetic and growth-related effects on foliar concentrations of total phenolics, PA and TNC in two different field studies. The first study contained nine different genotypes representing a range of genetic homogeneity, growing in a 2-year-old plantation on the coastal plain of North Carolina (NC), USA. The second study contained eight clones with different growth potentials planted in a 9-year-old clonal trial replicated at two sites (Georgia (GA) and South Carolina (SC), USA). In the first study (NC), we found no genetic effects on total phenolics, PA and TNC, and there was no relationship between genotype size and foliar biochemistry. In the second study, there were no differences in height growth between sites, but the SC site showed greater diameter (diameter at breast height (DBH)) and volume, most likely due to greater tree mortality (lower stocking) which reduced competition for resources and increased growth of remaining trees. We found a significant site × clone effect for total phenolics with lower productivity clones showing 27-30% higher total phenolic concentrations at the GA site where DBH and volume were lower. In contrast to the predictions of growth-defense theory, clone volume was positively associated with total phenolic concentrations at the higher volume SC site, and PA concentrations at the lower volume GA site. Overall, we found no evidence of a trade-off between genotype size and defense, and genetic potential for improved growth may include increased allocation to some secondary metabolites. These results imply that deployment of more productive loblolly pine genotypes will not reduce stand resistance to herbivory, but increased production of total phenolics and PA associated with higher genotype growth potential could reduce litter decomposition rates and therefore, nutrient availability.

Exposure to moderate concentrations of tropospheric ozone impairs tree stomatal response to carbon dioxide.

With rising concentrations of both atmospheric carbon dioxide (CO(2)) and tropospheric ozone (O(3)), it is important to better understand the interacting effects of these two trace gases on plant physiology affecting land-atmosphere gas exchange. We investigated the effect of growth under elevated CO(2) and O(3), singly and in combination, on the primary short-term stomatal response to CO(2) concentration in paper birch at the Aspen FACE experiment. Leaves from trees grown in elevated CO(2) and/or O(3) exhibited weaker short-term responses of stomatal conductance to both an increase and a decrease in CO(2) concentration from current ambient level. The impairement of the stomatal CO(2) response by O(3) most likely developed progressively over the growing season as assessed by sap flux measurements. Our results suggest that expectations of plant water-savings and reduced stomatal air pollution uptake under rising atmospheric CO(2) may not hold for northern hardwood forests under concurrently rising tropospheric O(3).

Leaf-level gas-exchange uniformity and photosynthetic capacity among loblolly pine (Pinus taeda L.) genotypes of contrasting inherent genetic variation.

Variation in leaf-level gas exchange among widely planted genetically improved loblolly pine (Pinus taeda L.) genotypes could impact stand-level water use, carbon assimilation, biomass production, C allocation, ecosystem sustainability and biogeochemical cycling under changing environmental conditions. We examined uniformity in leaf-level light-saturated photosynthesis (A(sat)), stomatal conductance (g(s)), and intrinsic water-use efficiency (A(sat)/g(s) or δ) among nine loblolly pine genotypes (selected individuals): three clones, three full-sib families and three half-sib families, during the early years of stand development (first 3 years), with each genetic group possessing varying amounts of inherent genetic variation. We also compared light- and CO(2)-response parameters between genotypes and examined the relationship between genotype productivity, gas exchange and photosynthetic capacity. Within full-sib, half-sib and clonal genotypes, the coefficient of variation (CV) for gas exchange showed no consistent pattern; the CV for g(s) and δ was similar within clonal (44.3-46.9 and 35.5-38.6%) and half-sib (41.0-49.3 and 36.8-40.9%) genotypes, while full-sibs showed somewhat higher CVs (46.9-56.0 and 40.1-45.4%). In contrast, the CVs for A(sat) were generally higher within clones. With the exception of δ, differences in gas exchange among genotypes were generally insignificant. Tree volume showed a significant positive correlation with A(sat) and δ, but the relationship varied by season. Individual-tree volume and genotype volume were positively correlated with needle dark respiration (R(d)). Our results suggest that uniformity in leaf-level physiological rates is not consistently related to the amount of genetic variation within a given genotype, and δ, A(sat) and R(d) were the leaf-level physiological parameters that were most consistently related to individual-tree and genotype productivity. An enhanced understanding of molecular and environmental factors that influence physiological variation within and between loblolly pine genotypes may improve assessments of genotype growth potential and sensitivity to global climate change.

Hydraulic redistribution of soil water by roots affects whole-stand evapotranspiration and net ecosystem carbon exchange.

*Hydraulic redistribution (HR) of water via roots from moist to drier portions of the soil occurs in many ecosystems, potentially influencing both water use and carbon assimilation. *By measuring soil water content, sap flow and eddy covariance, we investigated the temporal variability of HR in a loblolly pine (Pinus taeda) plantation during months of normal and below-normal precipitation, and examined its effects on tree transpiration, ecosystem water use and carbon exchange. *The occurrence of HR was explained by courses of reverse flow through roots. As the drought progressed, HR maintained soil moisture above 0.15 cm(3) cm(-3) and increased transpiration by 30-50%. HR accounted for 15-25% of measured total site water depletion seasonally, peaking at 1.05 mm d(-1). The understory species depended on water redistributed by the deep-rooted overstory pine trees for their early summer water supply. Modeling carbon flux showed that in the absence of HR, gross ecosystem productivity and net ecosystem exchange could be reduced by 750 and 400 g C m(-2) yr(-1), respectively. *Hydraulic redistribution mitigated the effects of soil drying on understory and stand evapotranspiration and had important implications for net primary productivity by maintaining this whole ecosystem as a carbon sink.

Decoupling the influence of leaf and root hydraulic conductances on stomatal conductance and its sensitivity to vapour pressure deficit as soil dries in a drained loblolly pine plantation.

The study examined the relationships between whole tree hydraulic conductance (K(tree)) and the conductance in roots (K(root)) and leaves (K(leaf)) in loblolly pine trees. In addition, the role of seasonal variations in K(root) and K(leaf) in mediating stomatal control of transpiration and its response to vapour pressure deficit (D) as soil-dried was studied. Compared to trunk and branches, roots and leaves had the highest loss of conductivity and contributed to more than 75% of the total tree hydraulic resistance. Drought altered the partitioning of the resistance between roots and leaves. As soil moisture dropped below 50%, relative extractable water (REW), K(root) declined faster than K(leaf). Although K(tree) depended on soil moisture, its dynamics was tempered by the elongation of current-year needles that significantly increased K(leaf) when REW was below 50%. After accounting for the effect of D on g(s), the seasonal decline in K(tree) caused a 35% decrease in g(s) and in its sensitivity to D, responses that were mainly driven by K(leaf) under high REW and by K(root) under low REW. We conclude that not only water stress but also leaf phenology affects the coordination between K(tree) and g(s) and the acclimation of trees to changing environmental conditions.

Soil respiration, root biomass, and root turnover following long-term exposure of northern forests to elevated atmospheric CO2 and tropospheric O3.

The Rhinelander free-air CO(2) enrichment (FACE) experiment is designed to understand ecosystem response to elevated atmospheric carbon dioxide (+CO(2)) and elevated tropospheric ozone (+O(3)). The objectives of this study were: to understand how soil respiration responded to the experimental treatments; to determine whether fine-root biomass was correlated to rates of soil respiration; and to measure rates of fine-root turnover in aspen (Populus tremuloides) forests and determine whether root turnover might be driving patterns in soil respiration. Soil respiration was measured, root biomass was determined, and estimates of root production, mortality and biomass turnover were made. Soil respiration was greatest in the +CO(2) and +CO(2) +O(3) treatments across all three plant communities. Soil respiration was correlated with increases in fine-root biomass. In the aspen community, annual fine-root production and mortality (g m(-2)) were positively affected by +O(3). After 10 yr of exposure, +CO(2) +O(3)-induced increases in belowground carbon allocation suggest that the positive effects of elevated CO(2) on belowground net primary productivity (NPP) may not be offset by negative effects of O(3). For the aspen community, fine-root biomass is actually stimulated by +O(3), and especially +CO(2) +O(3).

Forest response to elevated CO2 is conserved across a broad range of productivity.

Climate change predictions derived from coupled carbon-climate models are highly dependent on assumptions about feedbacks between the biosphere and atmosphere. One critical feedback occurs if C uptake by the biosphere increases in response to the fossil-fuel driven increase in atmospheric [CO(2)] ("CO(2) fertilization"), thereby slowing the rate of increase in atmospheric [CO(2)]. Carbon exchanges between the terrestrial biosphere and atmosphere are often first represented in models as net primary productivity (NPP). However, the contribution of CO(2) fertilization to the future global C cycle has been uncertain, especially in forest ecosystems that dominate global NPP, and models that include a feedback between terrestrial biosphere metabolism and atmospheric [CO(2)] are poorly constrained by experimental evidence. We analyzed the response of NPP to elevated CO(2) ( approximately 550 ppm) in four free-air CO(2) enrichment experiments in forest stands. We show that the response of forest NPP to elevated [CO(2)] is highly conserved across a broad range of productivity, with a stimulation at the median of 23 +/- 2%. At low leaf area indices, a large portion of the response was attributable to increased light absorption, but as leaf area indices increased, the response to elevated [CO(2)] was wholly caused by increased light-use efficiency. The surprising consistency of response across diverse sites provides a benchmark to evaluate predictions of ecosystem and global models and allows us now to focus on unresolved questions about carbon partitioning and retention, and spatial variation in NPP response caused by availability of other growth limiting resources.

Tropospheric O(3) compromises net primary production in young stands of trembling aspen, paper birch and sugar maple in response to elevated atmospheric CO(2).

Concentrations of atmospheric CO(2) and tropospheric ozone (O(3)) are rising concurrently in the atmosphere, with potentially antagonistic effects on forest net primary production (NPP) and implications for terrestrial carbon sequestration. Using free-air CO(2) enrichment (FACE) technology, we exposed north-temperate forest communities to concentrations of CO(2) and O(3) predicted for the year 2050 for the first 7 yr of stand development. Site-specific allometric equations were applied to annual nondestructive growth measurements to estimate above- and below-ground biomass and NPP for each year of the experiment. Relative to the control, elevated CO(2) increased total biomass 25, 45 and 60% in the aspen, aspen-birch and aspen-maple communities, respectively. Tropospheric O(3) caused 23, 13 and 14% reductions in total biomass relative to the control in the respective communities. Combined fumigation resulted in total biomass response of -7.8, +8.4 and +24.3% relative to the control in the aspen, aspen-birch and aspen-sugar maple communities, respectively. These results indicate that exposure to even moderate levels of O(3) significantly reduce the capacity of NPP to respond to elevated CO(2) in some forests.