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.

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.

A remotely sensed pigment index reveals photosynthetic phenology in evergreen conifers.

In evergreen conifers, where the foliage amount changes little with season, accurate detection of the underlying "photosynthetic phenology" from satellite remote sensing has been difficult, presenting challenges for global models of ecosystem carbon uptake. Here, we report a close correspondence between seasonally changing foliar pigment levels, expressed as chlorophyll/carotenoid ratios, and evergreen photosynthetic activity, leading to a "chlorophyll/carotenoid index" (CCI) that tracks evergreen photosynthesis at multiple spatial scales. When calculated from NASA's Moderate Resolution Imaging Spectroradiometer satellite sensor, the CCI closely follows the seasonal patterns of daily gross primary productivity of evergreen conifer stands measured by eddy covariance. This discovery provides a way of monitoring evergreen photosynthetic activity from optical remote sensing, and indicates an important regulatory role for carotenoid pigments in evergreen photosynthesis. Improved methods of monitoring photosynthesis from space can improve our understanding of the global carbon budget in a warming world of changing vegetation phenology.

Variability of sun-induced chlorophyll fluorescence according to stand age-related processes in a managed loblolly pine forest.

Leaf fluorescence can be used to track plant development and stress, and is considered the most direct measurement of photosynthetic activity available from remote sensing techniques. Red and far-red sun-induced chlorophyll fluorescence (SIF) maps were generated from high spatial resolution images collected with the HyPlant airborne spectrometer over even-aged loblolly pine plantations in North Carolina (United States). Canopy fluorescence yield (i.e., the fluorescence flux normalized by the light absorbed) in the red and far-red peaks was computed. This quantifies the fluorescence emission efficiencies that are more directly linked to canopy function compared to SIF radiances. Fluorescence fluxes and yields were investigated in relation to tree age to infer new insights on the potential of those measurements in better describing ecosystem processes. The results showed that red fluorescence yield varies with stand age. Young stands exhibited a nearly twofold higher red fluorescence yield than mature forest plantations, while the far-red fluorescence yield remained constant. We interpreted this finding in a context of photosynthetic stomatal limitation in aging loblolly pine stands. Current and future satellite missions provide global datasets of SIF at coarse spatial resolution, resulting in intrapixel mixture effects, which could be a confounding factor for fluorescence signal interpretation. To mitigate this effect, we propose a surrogate of the fluorescence yield, namely the Canopy Cover Fluorescence Index (CCFI) that accounts for the spatial variability in canopy structure by exploiting the vegetation fractional cover. It was found that spatial aggregation tended to mask the effective relationships, while the CCFI was still able to maintain this link. This study is a first attempt in interpreting the fluorescence variability in aging forest stands and it may open new perspectives in understanding long-term forest dynamics in response to future climatic conditions from remote sensing of SIF.

Assessing the interplay between canopy energy balance and photosynthesis with cellulose δ18O: large-scale patterns and independent ground-truthing.

There are few whole-canopy or ecosystem scale assessments of the interplay between canopy temperature and photosynthesis across both spatial and temporal scales. The stable oxygen isotope ratio (δ18O) of plant cellulose can be used to resolve a photosynthesis-weighted estimate of canopy temperature, but the method requires independent confirmation. We compare isotope-resolved canopy temperatures derived from multi-year homogenization of tree cellulose δ18O to canopy-air temperatures weighted by gross primary productivity (GPP) at multiple sites, ranging from warm temperate to boreal and subalpine forests. We also perform a sensitivity analysis for isotope-resolved canopy temperatures that showed errors in plant source water δ18O lead to the largest errors in canopy temperature estimation. The relationship between isotope-resolved canopy temperatures and GPP-weighted air temperatures was highly significant across sites (p < 0.0001, R2 = 0.82), thus offering confirmation of the isotope approach. The previously observed temperature invariance from temperate to boreal biomes was confirmed, but the greater elevation of canopy temperature above air temperature in the boreal forest was not. Based on the current analysis, we conclude that canopy temperatures in the boreal forest are as warm as those in temperate systems because day-time-growing-season air temperatures are similarly warm.

Soil-plant-atmosphere conditions regulating convective cloud formation above southeastern US pine plantations.

Loblolly pine trees (Pinus taeda L.) occupy more than 20% of the forested area in the southern United States, represent more than 50% of the standing pine volume in this region, and remove from the atmosphere about 500 g C m-2 per year through net ecosystem exchange. Hence, their significance as a major regional carbon sink can hardly be disputed. What is disputed is whether the proliferation of young plantations replacing old forest in the southern United States will alter key aspects of the hydrologic cycle, including convective rainfall, which is the focus of the present work. Ecosystem fluxes of sensible (Hs) and latent heat (LE) and large-scale, slowly evolving free atmospheric temperature and water vapor content are known to be first-order controls on the formation of convective clouds in the atmospheric boundary layer. These controlling processes are here described by a zero-order analytical model aimed at assessing how plantations of different ages may regulate the persistence and transition of the atmospheric system between cloudy and cloudless conditions. Using the analytical model together with field observations, the roles of ecosystem Hs and LE on convective cloud formation are explored relative to the entrainment of heat and moisture from the free atmosphere. Our results demonstrate that cloudy-cloudless regimes at the land surface are regulated by a nonlinear relation between the Bowen ratio Bo=Hs/LE and root-zone soil water content, suggesting that young/mature pines ecosystems have the ability to recirculate available water (through rainfall predisposition mechanisms). Such nonlinearity was not detected in a much older pine stand, suggesting a higher tolerance to drought but a limited control on boundary layer dynamics. These results enable the generation of hypotheses about the impacts on convective cloud formation driven by afforestation/deforestation and groundwater depletion projected to increase following increased human population in the southeastern United States.

Thermal optimality of net ecosystem exchange of carbon dioxide and underlying mechanisms.

• It is well established that individual organisms can acclimate and adapt to temperature to optimize their functioning. However, thermal optimization of ecosystems, as an assemblage of organisms, has not been examined at broad spatial and temporal scales. • Here, we compiled data from 169 globally distributed sites of eddy covariance and quantified the temperature response functions of net ecosystem exchange (NEE), an ecosystem-level property, to determine whether NEE shows thermal optimality and to explore the underlying mechanisms. • We found that the temperature response of NEE followed a peak curve, with the optimum temperature (corresponding to the maximum magnitude of NEE) being positively correlated with annual mean temperature over years and across sites. Shifts of the optimum temperature of NEE were mostly a result of temperature acclimation of gross primary productivity (upward shift of optimum temperature) rather than changes in the temperature sensitivity of ecosystem respiration. • Ecosystem-level thermal optimality is a newly revealed ecosystem property, presumably reflecting associated evolutionary adaptation of organisms within ecosystems, and has the potential to significantly regulate ecosystem-climate change feedbacks. The thermal optimality of NEE has implications for understanding fundamental properties of ecosystems in changing environments and benchmarking global models.

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.

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.

Substantial hysteresis in emergent temperature sensitivity of global wetland CH4 emissions.

Wetland methane (CH4) emissions ([Formula: see text]) are important in global carbon budgets and climate change assessments. Currently, [Formula: see text] projections rely on prescribed static temperature sensitivity that varies among biogeochemical models. Meta-analyses have proposed a consistent [Formula: see text] temperature dependence across spatial scales for use in models; however, site-level studies demonstrate that [Formula: see text] are often controlled by factors beyond temperature. Here, we evaluate the relationship between [Formula: see text] and temperature using observations from the FLUXNET-CH4 database. Measurements collected across the globe show substantial seasonal hysteresis between [Formula: see text] and temperature, suggesting larger [Formula: see text] sensitivity to temperature later in the frost-free season (about 77% of site-years). Results derived from a machine-learning model and several regression models highlight the importance of representing the large spatial and temporal variability within site-years and ecosystem types. Mechanistic advancements in biogeochemical model parameterization and detailed measurements in factors modulating CH4 production are thus needed to improve global CH4 budget assessments.

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.

Poplar plantation has the potential to alter the water balance in semiarid Inner Mongolia.

Poplar plantation is the most dominant broadleaf forest type in northern China. Since the mid-1990s plantation was intensified to combat desertification along China's northwestern border, i.e., within Inner Mongolia (IM). This evoked much concern regarding the ecological and environmental effects on areas that naturally grow grass or shrub vegetation. To highlight potential consequences of large-scale poplar plantations on the water budget within semiarid IM, we compared the growing season water balance (evapotranspiration (ET) and precipitation (PPT)) of a 3-yr old poplar plantation (Kp(3)) and a natural shrubland (Ks) in the Kubuqi Desert in western IM, and a 6-yr old poplar plantation (Bp(6)) growing under sub-humid climate near Beijing. The results showed that, despite 33% lower PPT at Kp(3), ET was 2% higher at Kp(3) (228 mm) as compared with Ks (223 mm) in May-September 2006. The difference derived mainly from higher ET at the plantation during drier periods of the growing season, which also indicated that the poplars must have partly transpired groundwater. Estimated growing season ET at Bp(6) was about 550 mm and more than 100% higher than at Kp(3). It is estimated that increases in leaf area index and net radiation at Kp(3) provide future potential for the poplars in Kubuqi to exceed the present ET and ET of the natural shrubland by 100-200%. These increases in ET are only possible through the permanent use of groundwater either directly by the trees or through increased irrigation. This may significantly change the water balance in the area (e.g., high ET at the cost of a reduction in the water table), which renders large-scale plantations a questionable tool in sustainable arid-land management.

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.

Drought during canopy development has lasting effect on annual carbon balance in a deciduous temperate forest.

* Climate change projections predict an intensifying hydrologic cycle and an increasing frequency of droughts, yet quantitative understanding of the effects on ecosystem carbon exchange remains limited. * Here, the effect of contrasting precipitation and soil moisture dynamics were evaluated on forest carbon exchange using 2 yr of eddy covariance and microclimate data from a 50-yr-old mixed oak woodland in northern Ohio, USA. * The stand accumulated 40% less carbon in a year with drought between bud-break and full leaf expansion (354 +/- 81 g C m(-2) yr(-1) in 2004 and 252 +/- 45 g C m(-2) yr(-1) in 2005). This was caused by greater suppression of gross ecosystem productivity (GEP; 16% = 200 g) than of ecosystem respiration (ER; 11% = 100 g) by drought. Suppressed GEP was traced to lower leaf area, lower apparent quantum yield and lower canopy conductance. The moisture sensitivity of ER may have been mediated by GEP. * The results highlight the vulnerability of the ecosystem to even a moderate drought, when it affects a critical aspect of development. Although the drought was preceded by rain, the storage capacity of the soil seemed limited to 1-2 wk, and therefore droughts longer than this are likely to impair productivity in the region.

Olevi Kull's lifetime contribution to ecology.

In this article dedicated to Olevi Kull (June 22, 1955-January 31, 2007), we draw on his writings (in English and translated) to outline his thoughts on the relationship between scientists and science. We provide a brief synthesis of his most important work, give a short account of his career and, to bring the man into focus, share some personal stories of interactions with him. Kull considered that for a personal understanding to become scientific knowledge it must be explained convincingly based on theory and empirical support, and then taught to others in both spoken and written words. He saw the last step as the main distinction between learning and science. Olevi Kull's approach to science relied on two principles: first, linking theory and experiments in challenging settings, e.g., to test the generality of his ideas he often challenged them in multi-layered, mixed-species canopies. Second, he insisted on setting experiments to test assumptions used in quantitative analyses or in explaining an observed outcome; this, at times, led to falsification of commonly held ideas, thus enhancing ecophysiological understanding. After describing Kull's application of these principles, we give a brief synthesis of his most important work, in which he demonstrated through experimentation and modeling how the vertical distribution of leaves in canopies is consistent with the acclimation of the photosynthetic apparatus. We also review some of his findings on the interactive effects of carbon dioxide and ozone on canopy photosynthesis.

Temperature responses to infrared-loading and water table manipulations in peatland mesocosms.

We initiated a multi-factor global change experiment to explore the effects of infrared heat loading (HT) and water table level (WL) treatment on soil temperature (T) in bog and fen peatland mesocosms. We found that the temperature varied highly by year, month, peatland type, soil depth, HT and WL manipulations. The highest effect of HT on the temperature at 25 cm depth was found in June for the bog mesocosms (3.34-4.27 degrees C) but in May for the fen mesocosms (2.32-4.33 degrees C) over the 2-year study period. The effects of WL in the bog mesocosms were only found between August and January, with the wet mesocosms warmer than the dry mesocosms by 0.48-2.03 degrees C over the 2-year study period. In contrast, wetter fen mesocosms were generally cooler by 0.16-3.87 degrees C. Seasonal changes of temperatures elevated by the HT also varied by depth and ecosystem type, with temperature differences at 5 cm and 10 cm depth showing smaller seasonal fluctuations than those at 25 cm and 40 cm in the bog mesocosms. However, increased HT did not always lead to warmer soil, especially in the fen mesocosms. Both HT and WL manipulations have also changed the length of the non-frozen season.

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.

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.