Foliar nitrogen concentrations and natural abundance of (15)N suggest nitrogen allocation patterns of Douglas-fir and mycorrhizal fungi during development in elevated carbon dioxide concentration and temperature

Pseudotsuga menziesii (Mirb.) Franco (Douglas-fir) seedlings were grown in a 2 x 2 factorial design in enclosed mesocosms at ambient temperature or 3.5 degrees C above ambient, and at ambient CO2 concentration ([CO2]) or 179 ppm above ambient. Two additional mesocosms were maintained as open controls. We measured the extent of mycorrhizal infection, foliar nitrogen (N) concentrations on both a weight basis (%N) and area basis (Narea), and foliar delta15N signatures (15N/14N ratios) from summer 1993 through summer 1997. Mycorrhizal fungi had colonized nearly all root tips across all treatments by spring 1994. Elevated [CO2] lowered foliar %N but did not affect N(area), whereas elevated temperature increased both foliar %N and Narea. Foliar delta15N was initially -1 per thousand and dropped by the final harvest to between -4 and -5 per thousand in the enclosed mesocosms, probably because of transfer of isotopically depleted N from mycorrhizal fungi. Based on the similarity in foliar delta15N among treatments, we conclude that mycorrhizal fungi had similar N allocation patterns across CO2 and temperature treatments. We combined isotopic and Narea data for 1993-94 to calculate fluxes of N for second- and third-year needles. Yearly N influxes were higher in second-year needles than in third-year needles (about 160 and 50% of initial leaf N, respectively), indicating greater sink strength in the younger needles. Influxes of N in second-year needles increased in response to elevated temperature, suggesting increased N supply from soil relative to plant N demands. In the elevated temperature treatments, N effluxes from third-year needles were higher in seedlings in elevated [CO2] than in ambient [CO2], probably because of increased N allocation below ground. We conclude that N allocation patterns shifted in response to the elevated temperature and [CO2] treatments in the seedlings but not in their fungal symbionts.     

Effects of elevated carbon dioxide concentration on growth and N2 fixation of young Robinia pseudoacacia

Effects of elevated CO2 concentration ([CO2]) on carbon (C) and nitrogen (N) uptake and N source partitioning (N2 fixation versus mineral soil N uptake) of 1-year-old Robinia pseudoacacia were determined in a dual 13C and 15N continuous labeling experiment. Seedlings were grown for 16 weeks in ambient (350 ppm) or elevated [CO2] (700 ppm) with 15NH4 15NO3 as the only mineral nitrogen source. Elevated [CO2] increased the fraction of new C in total C, but it did not alter C partitioning among plant compartments. Elevated [CO2] also increased the fraction of new N in total N and this was coupled with a shift in N source partitioning toward N2 fixation. Soil N uptake was unaffected by elevated [CO2], whereas N2 fixation was markedly increased by the elevated [CO2] treatment, mainly because of increased specific fixation (mg N mg(-1) nodule). As a result of increased N2 fixation, the C/N ratio of tree biomass tended to decrease in the elevated [CO2] treatment. Partitioning of N uptake among plant compartments was unaffected by elevated [CO2]. Total dry mass of root nodules doubled in response to elevated [CO2], but this effect was not significant because of the great variability of root nodule formation. Our results show that, in the N2-fixing R. pseudoacacia, increased C uptake in response to increased [CO2] is matched by increased N2 fixation, indicating that enhanced growth in elevated [CO2] might not be restricted by N limitations.     

Carbon assimilation and nitrogen in needles of fertilized and unfertilized field-grown Scots pine at natural and elevated concentrations of CO2

Effects of elevated CO2 concentration ([CO2]) on carbon assimilation and needle biochemistry of fertilized and unfertilized 25-30-year-old Scots pine (Pinus sylvestris L.) trees were studied in a branch bag experiment set up in a naturally regenerated stand. In each tree, one branch was enclosed in a bag supplied with ambient [CO2] (360 micromol mol(-1)), a second branch was enclosed in a bag supplied with elevated [CO2] (680 micromol(-1)) and a control branch was left unbagged. The CO2 treatments were applied from April 15 to September 15, starting in 1993 for unfertilized trees and in 1994 for fertilized trees, which were treated with N in June 1994. Net photosynthesis, amount and activity of Rubisco, N, starch, C:N ratio and SLA of needles were measured during the growing season of 1995. Light-saturated net photosynthetic rates of 1-year-old and current-year shoots measured at ambient [CO2] were not affected by growth [CO2] or N fertilization. Elevated [CO2] reduced the amount and activity of Rubisco, and the relative proportion of Rubisco to soluble proteins and N in needles of unfertilized trees. Elevated [CO2] also reduced the chlorophyll concentration (fresh weight basis) of needles of unfertilized trees. Soluble protein concentration of needles was not affected by growth [CO2]. Elevated [CO2] decreased the Rubisco:chlorophyll ratio in unfertilized and fertilized trees. Starch concentration was significantly increased at elevated [CO2] only in 1-year-old needles of fertilized trees. Elevated [CO2] reduced needle N concentration on a dry weight or structural basis (dry weight minus starch) in unfertilized trees, resulting in an increase in needle C:N ratio. Fertilization had no effect on soluble protein, chlorophyll, Rubisco or N concentration of needles. The decrease in the relative proportions of Rubisco and N concentration in needles of unfertilized trees at elevated [CO2] indicates reallocation of N resources away from Rubisco to nonphotosynthetic processes in other plant parts. Acclimation occurred in a single branch exposed to high [CO2], despite the large sink of the tree. The responses of 1-year-old and current-year needles to elevation of growth [CO2] were similar.     

Increased nitrogen-use efficiency of a short-rotation poplar plantation in elevated CO(2) concentration

We estimated nitrogen (N) use by trees of three poplar species exposed for 3 years to free air CO(2) enrichment (FACE) and determined whether the CO(2) treatment affected the future N availability of the plantation. Trees were harvested at the end of the first 3-year rotation and N concentration and content of woody tissues determined. Nitrogen uptake of fine roots and litter was measured throughout the first crop rotation. The results were related to previously published variations in soil N content during the same period. We estimated retranslocation from green leaves and processes determining N mobilization and immobilization, such as mineralization and nitrification, and N immobilization in litter and microbial biomass. In all species, elevated CO(2) concentration ([CO(2)]) significantly increased nitrogen-use efficiency (NUE; net primary productivity per unit of annual N uptake), decreased N concentration in most plant tissues, but did not significantly change cumulative N uptake by trees over the rotation. Total soil N was depleted more in elevated [CO(2)] than in ambient [CO(2)], although not significantly for all soil layers. The effect of elevated [CO(2)] was usually similar for all species, although differences among species were sometimes significant. During the first 3-year rotation, productivity of the plantation remained high in the elevated [CO(2)] treatment. However, we observed a potential reduction in N availability in response to elevated [CO(2)].     

Effects of elevated concentrations of atmospheric CO2 and tropospheric O3 on leaf litter production and chemistry in trembling aspen and paper birch communities

Human activities are increasing the concentrations of atmospheric carbon dioxide ([CO2]) and tropospheric ozone ([O3]), potentially leading to changes in the quantity and chemical quality of leaf litter inputs to forest soils. Because the quality and quantity of labile and recalcitrant carbon (C) compounds influence forest productivity through changes in soil organic matter content, characterizing changes in leaf litter in response to environmental change is critical to understanding the effects of global change on forests. We assessed the independent and combined effects of elevated [CO2] and elevated [O3] on foliar litter production and chemistry in aspen (Populus tremuloides Michx.) and birch-(Betula papyrifera Marsh.) aspen communities at the Aspen free-air CO2 enrichment (FACE) experiment in Rhinelander, WI. Litter was analyzed for concentrations of C, nitrogen (N), soluble sugars, lipids, lignin, cellulose, hemicellulose and C-based defensive compounds (soluble phenolics and condensed tannins). Concentrations of these chemical compounds in naturally senesced litter were similar in aspen and birch-aspen communities among treatments, except for N, the C:N ratio and lipids. Elevated [CO2] significantly increased C:N (+8.7%), lowered mean litter N concentration (-10.7%) but had no effect on the concentrations of soluble sugars, soluble phenolics and condensed tannins. Elevated [CO2] significantly increased litter biomass production (+33.3%), resulting in significant increases in fluxes of N, soluble sugars, soluble phenolics and condensed tannins to the soil. Elevated [O3] significantly increased litter concentrations of soluble sugars (+78.1%), soluble phenolics (+53.1%) and condensed tannins (+77.2%). There were no significant effects of elevated [CO2] or elevated [O3] on the concentrations of individual C structural carbohydrates (cellulose, hemicellulose and lignin). Elevated [CO2] significantly increased cellulose (+37.4%) input to soil, whereas elevated [O3] significantly reduced hemicellulose and lignin inputs to soil (-22.3 and -31.5%, respectively). The small changes in litter chemistry in response to elevated [CO2] and tropospheric [O3] that we observed, combined with changes in litter biomass production, could significantly alter the inputs of N, soluble sugars, condensed tannins, soluble phenolics, cellulose and lignin to forest soils in the future.     

Genotypic variation in physiological and growth responses of Populus tremuloides to elevated atmospheric CO2 concentration

Physiological and biomass responses of six genotypes of Populus tremuloides Michx., grown in ambient t (357 micromol mol(-1)) or twice ambient (707 micromol mol(-1)) CO2 concentration ([CO2]) and in low-N or high-N soils, were studied in 1995 and 1996 in northern Lower Michigan, USA. There was a significant CO2 x genotype interaction in photosynthetic responses. Net CO2 assimilation (A) was significantly enhanced by elevated [CO2] for five genotypes in high-N soil and for four genotypes in low-N soil. Enhancement of A by elevated [CO2] ranged from 14 to 68%. Genotypes also differed in their biomass responses to elevated [CO2], but biomass responses were poorly correlated with A responses. There was a correlation between magnitude of A enhancement by elevated [CO2] and stomatal sensitivity to CO2. Genotypes with low stomatal sensitivity to CO2 had a significantly higher A at elevated [CO2] than at ambient [CO2], but elevated [CO2] did not affect the ratio of intercellular [CO2] to leaf surface [CO2]. Stomatal conductance and A of different genotypes responded differentially to recovery from drought stress. Photosynthetic quantum yield and light compensation point were unaffected by elevated [CO2]. We conclude that P. tremuloides genotypes will respond differentially to rising atmospheric [CO2], with the degree of response dependent on other abiotic factors, such as soil N and water availability. The observed genotypic variation in growth could result in altered genotypic representation within natural populations and could affect the composition and structure of plant communities in a higher [CO2] environment in the future.     

Effects of elevated carbon dioxide concentration on growth and nitrogen fixation in Alnus glutinosa in a long-term field experiment

Nitrogen-fixing plant species may respond more positively to elevated atmospheric carbon dioxide concentrations ([CO2]) than other species because of their ability to maintain a high internal nutrient supply. A key factor in the growth response of trees to elevated [CO2] is the availability of nitrogen, although how elevated [CO2] influences the rate of N2-fixation of nodulated trees growing under field conditions is unclear. To elucidate this relationship, we measured total biomass, relative growth rate, net assimilation rate (NAR), leaf area and net photosynthetic rate of N2-fixing Alnus glutinosa (L.) Gaertn. (common alder) trees grown for 3 years in open-top chambers in the presence of either ambient or elevated atmospheric [CO2] and two soil N regimes: full nutrient solution or no fertilizer. Nitrogen fixation by Frankia spp. in the root nodules of unfertilized trees was assessed by the acetylene reduction method. We hypothesized that unfertilized trees would show similar positive growth and physiological responses to elevated [CO2] as the fertilized trees. Growth in elevated [CO2] stimulated (relative) net photosynthesis and (absolute) total biomass accumulation. Relative total biomass increased, and leaf nitrogen remained stable, only during the first year of the experiment. Toward the end of the experiment, signs of photosynthetic acclimation occurred, i.e., down-regulation of the photosynthetic apparatus. Relative growth rate was not significantly affected by elevated [CO2] because although NAR was increased, the effect on relative growth rate was negated by a reduction in leaf area ratio. Neither leaf area nor leaf P concentration was affected by growth in elevated [CO2]. Nodule mass increased on roots of unfertilized trees exposed to elevated [CO2] compared with fertilized trees exposed to ambient [CO2]. There was also a biologically significant, although not statistically significant, stimulation of nitrogenase activity in nodules exposed to elevated [CO2]. Root nodules of trees exposed to elevated [CO2] were smaller and more evenly spaced than root nodules of trees exposed to ambient [CO2]. The lack of an interaction between nutrient and [CO2] effects on growth, biomass and photosynthesis indicates that the unfertilized trees maintained similar CO2-induced growth and photosynthetic enhancements as the fertilized trees. This implies that alder trees growing in natural conditions, which are often limited by soil N availability, should nevertheless benefit from increasing atmospheric [CO2].     

A modeling analysis of the interaction between forest age and forest responsiveness to increasing CO2 concentration

Typically, forests have rotations of 10-200 years. On that time scale, anthropogenic increases in atmospheric carbon dioxide concentration ([CO2]) and the associated changes in climate are expected to be substantial. These changes will, therefore, almost certainly affect the growth of presently established forest stands. Most studies on the effects of increasing [CO2] on tree growth have been made with young plants. However, the growth of trees within a forest stand varies with age. As a consequence, it is difficult to infer from the available experimental data how rising [CO2] will affect forest productivity over a full rotation. In this study, various mechanisms that may account for the slowing of forest growth with age were introduced into the forest growth model CenW, to assess how these processes affect the modeled growth response to increasing [CO2]. Inclusion of allocation shifts with tree height, individual tree mortality, changing respiration load and nutrient dynamics or age-based reductions in photosynthetic capacity had only small effects on the response to increasing [CO2]. However, when photosynthesis of mature trees was decreased as a function of size, then the growth response to increasing [CO2] was reduced because, at the same age, trees were larger in elevated than in ambient [CO2]. No simple and generally valid interactions between increasing [CO2] and forest age were identified because of the large number of interacting processes, all of which are incompletely understood. Important age x climate change interactions on productivity must occur in real forests and need to be considered to understand likely future trends. However, these interactions are complex and difficult to test. It is therefore not yet possible to predict with confidence the modification of the CO2 response by forest age.     

Effects of elevated concentrations of atmospheric CO2 and tropospheric O3 on decomposition of fine roots

Rising atmospheric carbon dioxide (CO2) concentration ([CO2]) could alter terrestrial carbon (C) cycling by affecting plant growth, litter chemistry and decomposition. How the concurrent increase in tropospheric ozone (O3) concentration ([O3]) will interact with rising atmospheric [CO2] to affect C cycling is unknown. A major component of carbon cycling in forests is fine root production, mortality and decomposition. To better understand the effects of elevated [CO2] and [O3] on the dynamics of fine root C, we conducted a combined field and laboratory incubation experiment to monitor decomposition dynamics and changes in fine root litter chemistry. Free-air CO2 enrichment (FACE) technology at the FACTS-II Aspen FACE project in Rhinelander, Wisconsin, elevated [CO2] (535 microl 1-1) and [O3] (53 nl 1-1) in intact stands of pure trembling aspen (Populus tremuloides Michx.) and in mixed stands of trembling aspen plus paper birch (Betula papyrifera Marsh.) and trembling aspen plus sugar maple (Acer saccharum Marsh.). We hypothesized that the trees would react to increased C availability (elevated [CO2]) by increasing allocation to C-based secondary compounds (CBSCs), thereby decreasing rates of decomposition. Because of its lower growth potential, we reasoned this effect would be greatest in the aspen-maple community relative to the aspen and aspen-birch communities. As a result of decreased C availability, we expected elevated [O3] to counteract shifts in C allocation induced by elevated [CO2]. Concentrations of CBSCs were rarely significantly affected by the CO2 and O3 treatments in decomposing fine roots. Rates of microbial respiration and mass loss from fine roots were unaffected by the treatments, although the production of dissolved organic C differed among communities. We conclude that elevated [CO2] and [O3] induce only small changes in fine root chemistry that are insufficient to significantly influence fine root decomposition. If changes in soil C cycling occur in the future, they will most likely be brought about by changes in litter production.     

Ammonium and nitrate acquisition by plants in response to elevated CO2 concentration: the roles of root physiology and architecture

We examined changes in root system architecture and physiology and whole-plant patterns of nitrate reductase (NR) activity in response to atmospheric CO2 enrichment and N source to determine how changes in the form of N supplied to plants interact with rising CO2 concentration ([CO2]). Seedlings of Betula alleghaniensis Britt. and Pinus strobus L., which differ in growth rate, root architecture, and the partitioning of NR activity between leaves (Betula) and roots (Pinus), were grown in ambient (400 microl l(-1)) and elevated (800 microl l(-1)) [CO2] and supplied with either nitrate (NO3-) or ammonium (NH4+) as their sole N source. After 15 weeks of growth, plants were harvested and root system architecture, N uptake kinetics, and NR activity measured. Betula alleghaniensis responded to elevated [CO2] with significant increases in growth, regardless of the source of N. Pinus strobus showed no significant response in biomass production or allocation to elevated [CO2]. Both species exhibited significantly greater growth with NH4+ than with NO3-, along with lower root:shoot biomass ratios. Betula showed significant increases in total root length in response to elevated [CO2]. However, root N uptake rates in Betula (for both NO3- and NH4+) were either reduced or unchanged by elevated [CO2]. Pinus showed the opposite response to elevated [CO2], with no change in root architecture, but an increase in maximal uptake rates in response to elevated [CO2]. Nitrate reductase activity (on a mass basis) was reduced in leaves of Betula in elevated [CO2], but did not change in other tissues. Nitrate reductase activity was unaffected by elevated [CO2] in Pinus. Scaling this response to the whole-plant, NR activity was reduced in elevated [CO2] in Betula but not in Pinus. However, because Betula plants were larger in elevated [CO2], total whole-plant NR activity was unaffected.     

Temperature responses of growth and wood anatomy in European beech saplings grown in different carbon dioxide concentrations

Effects of temperature on growth and wood anatomy were studied in young European beech (Fagus sylvatica L.) grown in 7-l pots for 2.5 years in field-phytotron chambers supplied with an ambient (approximately 400 micromol mol-1) or elevated (approximately 700 micromol mol-1) carbon dioxide concentration ([CO2]). Temperatures in the chambers ranged in increments of 2 degrees C from -4 degrees C to +4 degrees C relative to the long-term mean monthly (day and night) air temperature in Berlin-Dahlem. Soil was not fertilized and soil water and air humidity were kept constant. Data were evaluated by regression analysis. At final harvest, stem diameter was significantly greater at increased temperature (Delta1 degrees C: 2.4%), stems were taller (Delta1 degrees C: 8.5%) and stem mass tree-1 (Delta1 degrees C: 10.9%) and leaf area tree-1(Delta1 degrees C: 6.5%) were greater. Allocation pattern was slightly influenced by temperature: leaf mass ratio and leaf area ratio decreased with increasing temperature (Delta1 degrees C: 2.3% and 2.2% respectively), whereas stem mass/total mass increased (Delta1 degrees C: 2.1%). Elevated [CO2] enhanced height growth by 8.8% and decreased coarse root mass/total mass by 10.3% and root/shoot ratio by 11.7%. Additional carbon was mainly invested in aboveground growth. At final harvest a synergistic interaction between elevated [CO2] and temperature yielded trees that were 3.2% taller at -4 degrees C and 12.7% taller at +4 degrees C than trees in ambient [CO2]. After 2.5 seasons, cross-sectional area of the oldest stem part was approximately 32% greater in the +4 degrees C treatment than in the -4 degrees C treatment, and in the last year approximately 67% more leaf area/unit tree ring area was produced in the highest temperature regime compared with the lowest. Elevated [CO2] decreased mean vessel area of the 120 largest vessels per mm2 by 5.8%, causing a decrease in water conducting capacity. There was a positive interaction between temperature and elevated [CO2] for relative vessel area, which was approximately 38% higher at +4 degrees C than at -4 degrees C in elevated [CO2] compared with ambient [CO2]. Overall, temperature had a greater effect on growth than [CO2], but elevated [CO2] caused quantitative changes in wood anatomy.     

Fine root respiration in mature eastern white pine (Pinus strobus) in situ: the importance of CO(2) in controlled environments

We measured seasonal fine root respiration rate in situ while controlling chamber temperature and [CO(2)]. Atmospheric [CO(2)] ([CO(2)](a)) and measured soil [CO(2)] ([CO(2)](s)) were alternately delivered to a cuvette containing intact fine roots of eastern white pine (Pinus strobus L.). Respiration rates were consistently higher in [CO(2)](a) than in [CO(2)](s) and were almost three times higher during midsummer. Respiration rates were immediately reversed after returning to the alternate [CO(2)] (i.e., [CO(2)](a) --> [CO(2)](s) --> [CO(2)](a), and vice versa) suggesting a direct effect of elevated [CO(2)] on apparent respiration. Soil-[CO(2)]-based respiration rates decreased with increasing [CO(2)] on a dry mass and tissue [N] basis. We conclude that estimates of soil CO(2) flux and soil carbon budgets may be improved by more completely accounting for the rhizosphere microclimate (i.e., soil temperature and [CO(2)](s)) during measurement of fine root respiration.     

Photosynthetic acclimation of overstory Populus tremuloides and understory Acer saccharum to elevated atmospheric CO2 concentration: interactions with shade and soil nitrogen

We exposed Populus tremuloides Michx. and Acer saccharum Marsh. to a factorial combination of ambient and elevated atmospheric CO2 concentrations ([CO2]) and high-nitrogen (N) and low-N soil treatments in open-top chambers for 3 years. Our objective was to compare photosynthetic acclimation to elevated [CO2] between species of contrasting shade tolerance, and to determine if soil N or shading modify the acclimation response. Sun and shade leaf responses to elevated [CO2] and soil N were compared between upper and lower canopy leaves of P. tremuloides and between A. saccharum seedlings grown with and without shading by P. tremuloides. Both species had higher leaf N concentrations and photosynthetic rates in high-N soil than in low-N soil, and these characteristics were higher for P. tremuloides than for A. saccharum. Electron transport capacity (Jmax) and carboxylation capacity (Vcmax) generally decreased with atmospheric CO2 enrichment in all 3 years of the experiment, but there was no evidence that elevated [CO2] altered the relationship between them. On a leaf area basis, both Jmax and Vcmax acclimated to elevated [CO2] more strongly in shade leaves than in sun leaves of P. tremuloides. However, the apparent [CO2] x shade interaction was largely driven by differences in specific leaf area (m2 g-1) between sun and shade leaves. In A. saccharum, photosynthesis acclimated more strongly to elevated [CO2] in sun leaves than in shade leaves on both leaf area and mass bases. We conclude that trees rooted freely in the ground can exhibit photosynthetic acclimation to elevated [CO2], and the response may be modified by light environment. The hypothesis that photosynthesis acclimates more completely to elevated [CO2] in shade-tolerant species than in shade-intolerant species was not supported.     

Atmospheric carbon dioxide concentration, nitrogen availability, temperature and the photosynthetic capacity of current-year Norway spruce shoots

Branches of field-grown Norway spruce (Picea abies (L.) Karst.) trees were exposed to either long-term ambient or to elevated CO2 concentrations ([CO2]) using the branch bag technique. The light-saturated photosynthetic rates (A(max)) of current-year shoots differing in nitrogen (N) status were measured at various temperatures and at either ambient (360 micromol mol(-1), AMB) or elevated (ambient + 350 micromol mol(-1), EL) [CO2]. The value of A(max) was determined at various intercellular [CO2]s (A/Ci curves) and used to normalize photosynthetic rates to the mean treatment C(i) values, which were 200 micromol mol(-1) (AMB) and 450 micromol mol(-1) (EL), respectively. Needle N status and temperature strongly affected A(max). The response to N increased with temperature, and the photosynthetic temperature optimum increased with N status. This was assumed to be a result of reduced mesophyll CO2 conductance. The relative increase of Amax in the EL treatment compared to the AMB treatment varied from 15 to 90%, and increased with temperature, but decreased with N status. Nevertheless, the absolute photosynthetic response to EL increased with shoot N status. The relative increase in the instantaneous response of A(max) to elevated [CO2] was about 20% higher than the long-term response, i.e., there was downward acclimation in Amax in response to elevated [CO2]. The photosynthetic temperature optimum increased 4 degrees C with either a short- or a long-term increase in [CO2]. The bag treatment itself increased A(max) by approximately 16% and the temperature optimum of A(max) by approximately 3 degrees C.     

Leaf photosynthetic characteristics of silver birch during three years of exposure to elevated concentrations of CO2 and O3 in the field

Effects of elevated concentrations of carbon dioxide ([CO2]) and ozone ([O3]) on photosynthesis and related biochemistry of two European silver birch (Betula pendula Roth) clones were studied under field conditions during 1999-2001. Seven-year-old trees of Clones 4 and 80 were exposed for 3 years to the following treatments in an open-top chamber experiment: outside control (OC), chamber control (CC), 2x ambient [CO2] (EC), 2x ambient [O3] (EO) and 2x ambient [CO2] + 2x ambient [O3] (EC+EO). During the experiment, gas exchange, chlorophyll fluorescence, amount and activity of Rubisco, concentrations of chlorophyll, soluble protein, soluble sugars, starch, nitrogen (N) and carbon:nitrogen (C:N) ratio were determined in short- and long-shoot leaves. Elevated [CO2] increased photosynthetic rate by around 30% when measurements were made at the growth [CO2]. When measured at ambient [CO2], photosynthesis was around 15% lower in EC trees than in CC trees. This was related to a approximately 10% decrease in total leaf N, to 26 and 20% decreases in the amount and activity of Rubisco, respectively, and to a 49% increase in starch concentration in elevated [CO2]. Elevated [O3] had no significant effect on gas exchange parameters and its effect on biochemistry was small in both clones. However, elevated [O3] decreased the proportion of Rubisco in total soluble proteins and the apparent quantum yield of photosystem II (PSII) photochemistry in light and increased non-photochemical quenching in 2000. The interactive effect of CO2 and O3 was variable. Elevated [O3] decreased chlorophyll concentration only in EO trees, and the EC+EO treatment decreased the total activity of Rubisco and increased the C:N ratio more than the EO treatment alone. The small effect of elevated [O3] on photosynthesis indicates that these young silver birches were fairly tolerant to annual [O3] exposures that were 2-3 times higher than the AOT40 value of 10 ppm.h. set as a critical dose for forest trees.     

Microelectrode technique for in situ measurement of carbon dioxide concentrations in xylem sap of trees

We developed a new microelectrode technique for measuring CO2 concentration ([CO2]) in xylem sap of trees. This technique enabled us to make rapid and continuous measurements of xylem sap [CO2] in situ. In this report, we discuss the methodology and establish the feasibility of the technique. We also describe calibration procedures, temperature sensitivity, field use and other characteristics of the microelectrodes. An example of data collected in the field is provided. Microelectrode calibration was accomplished at constant temperature in air of known [CO2]. When sampling temperature differed from calibration temperature, correction was necessary. We developed an equation to correct for temperatures between 15 and 35 degrees C when calibration was conducted at 25 degrees C. Equations based on Henry's Law were used to convert measured gas phase [CO2] (%) to concentration of all products of CO2 dissolved in sap (mmol l(-1)). We inserted microelectrodes into stems of three tree species to measure diurnal changes in [CO2] in the xylem sap. A diurnal pattern with depression during the day and elevation at night was observed. Mean daily [CO2] ranged from 1.6 to 10.3 mmol l(-1). Microelectrodes were suitable for making diurnal measurements for up to 7 days without recalibration. We also used the microelectrodes to measure [CO2] of soil in situ. Soil [CO2] ranged from 1 to 4% (gas phase), with little diurnal variation.     

Interactive effects of elevated CO2 concentration and nitrogen supply on partitioning of newly fixed 13C and 15N between shoot and roots of pedunculate oak seedlings (Quercus robur)

Pedunculate oak (Quercus robur L.) seedlings were grown for 3 or 4 months (second- and third-flush stages) in greenhouses at two atmospheric CO2 concentrations ([CO2]) (350 or 700 micromol mol(-1)) and two nitrogen fertilization regimes (6.1 or 0.61 mmol N l(-1) nutrient solution). Combined effects of [CO2] and nitrogen fertilization on partitioning of newly acquired carbon (C) and nitrogen (N) were assessed by dual 13C and 15N short-term labeling of seedlings at the second- or third-flush stage of development. In the low-N treatment, root growth, but not shoot growth, was stimulated by elevated [CO2], with the result that shoot/root biomass ratio declined. At the second-flush stage, overall seedling biomass growth was increased (13%) by elevated [CO2] regardless of N fertilization. At the third-flush stage, elevated [CO2] increased growth sharply (139%) in the high-N but not the low-N treatment. Root/shoot biomass ratios were threefold higher in the low-N treatment relative to the high-N treatment. At the second-flush stage, leaf area was 45-51% greater in the high-N treatment than in the low-N treatment. At the-third flush stage, there was a positive interaction between the effects of N fertilization and [CO2] on leaf area, which was 93% greater in the high-N/elevated [CO2] treatment than in the low-N/ambient [CO2] treatment. Specific leaf area was reduced (17-25%) by elevated [CO2], whereas C and N concentrations of seedlings increased significantly in response to either elevated [CO2] or high-N fertilization. At the third-flush stage, acquisition of C and N per unit dry mass of leaf and fine root was 51 and 77% greater, respectively, in the elevated [CO2]/high-N fertilization treatment than in the ambient [CO2]/low-N fertilization treatment. However, there was dilution of leaf N in response to elevated [CO2]. Partitioning of newly acquired C and N between shoot and roots was altered by N fertilization but not [CO2]. More newly acquired C and N were partitioned to roots in the low-N treatment than in the high-N treatment.     

Leaf respiration at different canopy positions in sweetgum (Liquidambar styraciflua) grown in ambient and elevated concentrations of carbon dioxide in the field

Trees exposed to elevated CO2 partial pressure ([CO2]) generally show increased rates of photosynthesis and growth, but effects on leaf respiration are more variable. The causes of this variable response are unresolved. We grew 12-year-old sweetgum trees (Liquidambar styraciflua L.) in a Free-Air CO2 Enrichment (FACE) facility in ambient [CO2] (37/44 Pa daytime/nighttime) and elevated [CO2] (57/65 Pa daytime/nighttime) in native soil at Oak Ridge National Environmental Research Park. Nighttime respiration (R(N)) was measured on leaves in the upper and lower canopy in the second (1999) and third (2000) growing seasons of CO2 fumigation. Leaf respiration in the light (R(L)) was estimated by the technique of Brooks and Farquhar (1985) in the upper canopy during the third growing season. There were no significant short-term effects of elevated [CO2] on R(N) or long-term effects on R(N) or R(L), when expressed on an area, mass or nitrogen (N) basis. Upper-canopy leaves had 54% higher R(N) (area basis) than lower-canopy leaves, but this relationship was unaffected by CO2 growth treatment. In August 2000, R(L) was about 40% of R(N) in the upper canopy. Elevated [CO(2)] significantly increased the number of leaf mitochondria (62%), leaf mass per unit area (LMA; 9%), and leaf starch (31%) compared with leaves in ambient [CO(2)]. Upper-canopy leaves had a significantly higher number of mitochondria (73%), N (53%), LMA (38%), sugar (117%) and starch (23%) than lower-canopy leaves. Growth in elevated [CO2] did not affect the relationships (i.e., intercept and slope) between R(N) and the measured leaf characteristics. Although no factor explained more than 45% of the variation in R(N), leaf N and LMA were the best predictors for R(N). Therefore, the response of RN to CO2 treatment and canopy position was largely dependent on the magnitude of the effect of elevated [CO2] or canopy position on these characteristics. Because elevated [CO2] had little or no effect on N or LMA, there was no effect on R(N). Canopy position had large effects on these leaf characteristics, however, such that upper-canopy leaves exhibited higher R(N) than lower-canopy leaves. We conclude that elevated [CO2] does not directly impact leaf respiration in sweetgum and that barring changes in leaf nitrogen or leaf chemical composition, long-term effects of elevated [CO2] on respiration in this species will be minimal.     

Influence of elevated CO2 and mycorrhizae on nitrogen acquisition: contrasting responses in Pinus taeda and Liquidambar styraciflua

An understanding of root system capacity to acquire nitrogen (N) is critical in assessing the long-term growth impact of rising atmospheric CO2 concentration ([CO2]) on trees and forest ecosystems. We examined the effects of mycorrhizal inoculation and elevated [CO2] on root ammonium (NH4+) and nitrate (NO3-) uptake capacity in sweetgum (Liquidambar styraciflua L.) and loblolly pine (Pinus taeda L.). Mycorrhizal treatments included inoculation of seedlings with the arbuscular mycorrhizal (AM) fungus Glomus intraradices Schenck & Smith in sweetgum and the ectomycorrhizal (EM) fungus Laccaria bicolor (Maire) Orton in loblolly pine. These plants were then equally divided between ambient and elevated [CO2] treatments. After 6 months of treatment, root systems of both species exhibited a greater uptake capacity for NH4+ than for NO3-. In both species, mycorrhizal inoculation significantly increased uptake capacity for NO3-, but not for NH4+. In sweetgum, the mycorrhizal effect on NO3- and NH4+ uptake capacity depended on growth [C02]. Similarly, in loblolly pine, the mycorrhizal effect on NO3- uptake capacity depended on growth [CO2], but the effect on NH4+ uptake capacity did not. Mycorrhizal inoculation significantly enhanced root nitrate reductase activity (NRA) in both species, but elevated [CO2] increased root NRA only in sweetgum. Leaf NRA in sweetgum did not change significantly with mycorrhizal inoculation, but increased in response to [CO2]. Leaf NRA in loblolly pine was unaffected by either treatment. The results indicate that the mycorrhizal effect on specific root N uptake in these species depends on both the form of inorganic N and the mycorrhizal type. However, our data show that in addressing N status of plants under high [CO2], reliable prediction is possible only when information about other root system adjustments (e.g., biomass allocation to fine roots) is simultaneously considered.     

Advances of study on atmospheric methane oxidation （consumption） in forest soil

Next to CO2, methane (CH4) is the second important contributor to global warming in the atmosphere and global atmospheric CH4 budget depends on both CH4 sources and sinks. Unsaturated soil is known as a unique sink for atmospheric CH4 in terrestrial ecosystem. Many comparison studies proved that forest soil had the biggest capacity of oxidizing atmospheric CH4 in various unsaturated soils. However, up to now, there is not an overall review in the aspect of atmospheric CH4 oxidation (consumption) in forest soil. This paper analyzed advances of studies on the mechanism of atmospheric CH4 oxidation, and related natural factors (Soil physical and chemical characters, temperature and moisture, ambient main greenhouse gases concentrations, tree species, and forest fire) and anthropogenic factors (forest clear-cutting and thinning, fertilization, exogenou saluminum salts and atmospheric deposition, adding biocides, and switch of forest land use) in forest soils. It was believed that OH4 consumption rate by forest soil was limited by diffusion and sensitive to changes in water status and temperature of soil.CH4 oxidation was also particularly sensitive to soil C/N, Ambient CO2, CH4 and N2O concentrations, tree species and forest fire.In most cases, anthropogenic disturbances will decrease atmospheric CH4 oxidation, thus resulting in the elevating of atmos-pheric CH4. Finally, the author pointed out that our knowledge of atmospheric CH4 oxidation (consumption) in forest soil was insufficient. In order to evaluate the contribution of forest soils to atmospheric CH4 oxidation and the role of forest played in the process of global environmental change, and to forecast the trends of global warming exactly, more researchers need to studiesfurther on CH4 oxidation in various forest soils of different areas.