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.     

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.     

Stomatal response ofPinus sylvestriformis to elevated CO2 concentrations during the four years of exposure

Four-year-oldPinus sylvestriformis were exposed for four growing seasons in open top chambers to ambient CO₂ concentration (approx. 350 μmol·mol⁻¹) and high CO₂ concentrations (500 and 700 μmol·mol⁻¹) at Research Station of Changbai Mountain Forest Ecosystems, Chinese Academy of Sciences at Antu Town, Jilin Province, China (42°N, 128°E). Stomatal response to elevated CO₂ concentrations was examined by stomatal conductance (g _s), ratio of intercellular to ambient CO₂ concentration (c _i/c _a) and stomatal number. Reciprocal transfer experiments of stomatal conductance showed that stomatal conductance in high-[CO₂]-grown plants increased in comparison with ambient-[CO₂]-grown plants when measured at their respective growth CO₂ concentration and at the same measurement CO₂ concentration (except a reduction in 700 μmol·mol⁻¹ CO₂. grown plants compared with plants on unchambered field when measured at growth CO₂ concentration and 350 μmol·mol⁻¹CO₂). High-[CO₂]-grown plants exhibited lowerc _i/c _a ratios than ambient-[CO₂]-grown plants when measured at their respective growth CO₂ concentration. However,c _i/c _a ratios increased for plants grown in high CO₂ concentrations compared with control plants when measured at the same CO₂ concentration. There was no significant difference in stomatal number per unit long needle between elevated and ambient CO₂. However, elevated CO₂ concentrations reduced the total stomatal number of whole needle by the decline of stomatal line and changed the allocation pattern of stomata between upper and lower surface of needle. Foundation Item: This research was supported by National Basic Research Program of China (2002CB412502), Project of Key program of the National Natural Science Foundation of China (90411020) and National Natural Science Foundation of China (30400051). Biography: ZHOU Yu-mei (1973-), female, Ph. Doctor, assistant research fellow, Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang 110016, P. R. China. Responsible editor: Song Funan     

四个生长季高浓度CO_2处理长白松的气孔响应(英文)

在长白山站以开顶箱方式对4 年生长白松连续4 个生长季进行CO2 处理,包括700 和500 μmolmol-1 高浓度CO2,以及接受空气CO2的对照箱和不扣箱的裸露地条件(约350 μmolmol-1 CO2),通过测定气孔导度(gs),ci/ca比及气孔数量等指标评价气孔对高浓度CO2的响应。气孔导度及ci/ca比的转换实验表明,在各自生长CO2下和在相同测定CO2 下进行比较时,生长在高浓度CO2 下植株的气孔导度要高于空气CO2 下对照组植株的气孔导度(除700 μmol mol-1 CO2 下的植株在生长CO2 浓度下及在350 μmol mol-1 CO2 下测定时的气孔导度低于裸地植株外)。在各自生长CO2浓度下测定时,高浓度CO2下植株的ci/ca比低于对照组植株,但在相同测定CO2浓度下比较时,却是高浓度CO2下植株的ci/ca高于对照组植株的ci/ca比。高浓度CO2下植株与对照组植株在每单位长度气孔数量上无明显差异,但高浓度CO2通过降低气孔线数使长白松当年生针叶的总气孔数量降低,并且改变了气孔在针叶上、下表面的分配模式。表4 参18。     

Propagation of trembling aspen and hybrid poplar for agroforestry: potential benefits of elevated CO2 in the greenhouse

We examined the usefulness of elevated CO₂ in the greenhouse to aid in early selection of genotypes and in the propagation of Populus tremuloides Michaux (aspen) and hybrid poplars for agroforestry, afforestation, or reclamation. Growth in elevated (800 ppm) vs ambient (375 ppm) CO₂ for 95 days resulted in greater height (14%), stem caliper (16%), overall biomass, and proportional allocation of biomass to roots as well as elevated net assimilation and water-use-efficiency. Aspen clones selected for superior growth (based on phenotypic selection) broke bud significantly earlier than unselected clones under both CO₂ levels; superior clones were also taller with greater stem caliper under both CO₂ treatments (but not significant). Under ambient CO₂ male aspen were taller than females while under elevated CO₂ female aspen were taller than males and also had greater caliper (but not significant). Hybrid poplar grown under elevated (vs ambient) CO₂ broke bud significantly earlier and had significantly greater net assimilation and water-use efficiency; they were also taller with greater caliper (but not significant). Differences in performance among the eight hybrid clones were enhanced by growth under elevated CO₂. Under ambient CO₂, P38P38 was the clone that broke bud earliest, was tallest with greater caliper and had a lower shoot:root biomass ratio. Sargentii, on the other hand, was the shortest with the smallest caliper. The implications of these results need to be considered in the context of using aspen and hybrid poplar for large-scale agroforestry, afforestation and reclamation across Canada. This revised version was published online in August 2006 with corrections to the Cover Date.     

Respiratory responses of Scots pine stems to 5 years of exposure to elevated CO2 concentration and temperature

Stem respiration in 20-year-old Scots pine (Pinus sylvestris L.) trees was examined following 5 years of exposure to ambient conditions (CON), elevated atmospheric carbon dioxide concentration ([CO2]) (ambient + 350 micromol mol(-1), (EC)), elevated temperature (ambient + 2-6 degrees C, (ET)) or a combination of elevated [CO2] and elevated temperature (ECT). Stem respiration varied seasonally regardless of the treatment and displayed a similar trend to temperature, with maximum rates occurring around Day 190 in summer and minimum rates in winter. Respiration normalized to 15 degrees C (R15) was higher in the growing season than in the non-growing season, whereas the temperature coefficient (Q10) was lower in the growing season. Annually averaged R15 was 0.36, 0.43, 0.40 and 0.44 micromol m(-2) s(-1) under CON, EC, ET and ECT conditions, respectively, whereas the corresponding values for total stem respiration were 6.55, 7.69, 7.50 and 7.90 mol m(-2) year(-1). The EC, ET and ECT treatments increased R15 by 18, 11 and 22%, respectively, relative to CON, and increased the modeled annual total stem respiration by 18, 15 and 21%. The increase in modeled annual stem respiration under EC and ECT conditions was caused mainly by higher maintenance respiration (22 and 25%, respectively, whereas the increase in growth respiration was 9 and 12%). Growth respiration was unaltered by ET. The treatments did not significantly affect the respiratory response to stem temperature; the mean Q10 value was 2.04, 2.10, 1.99 and 2.12 in the CON, EC, ET and ECT treatments, respectively. It is suggested that the increase in stem respiration was partly a result of the increased growth rate. We conclude that elevated [CO2] increased the maintenance component of respiration more than the growth component.     

Carbon gain and bud physiology in Populus tremuloides and Betula papyrifera grown under long-term exposure to elevated concentrations of CO2 and O3

Paper birch (Betula papyrifera Marsh.) and three trembling aspen clones (Populus tremuloides Michx.) were studied to determine if alterations in carbon gain in response to an elevated concentration of CO(2) ([CO(2)]) or O(3) ([O(3)]) or a combination of both affected bud size and carbohydrate composition in autumn, and early leaf development in the following spring. The trees were measured for gas exchange, leaf size, date of leaf abscission, size and biochemical characteristics of the overwintering buds and early leaf development during the 8th-9th year of free-air CO(2) and O(3) exposure at the Aspen FACE site located near Rhinelander, WI. Net photosynthesis was enhanced 49-73% by elevated [CO(2)], and decreased 13-30% by elevated [O(3)]. Elevated [CO(2)] delayed, and elevated [O(3)] tended to accelerate, leaf abscission in autumn. Elevated [CO(2)] increased the ratio of monosaccharides to di- and oligosaccharides in aspen buds, which may indicate a lag in cold acclimation. The total carbon concentration in overwintering buds was unaffected by the treatments, although elevated [O(3)] decreased the amount of starch by 16% in birch buds, and reduced the size of aspen buds, which may be related to the delayed leaf development in aspen during the spring. Elevated [CO(2)] generally ameliorated the effects of elevated [O(3)]. Our results show that both elevated [CO(2)] and elevated [O(3)] have the potential to alter carbon metabolism of overwintering buds. These changes may cause carry-over effects during the next growing season.     

Leaf photosynthetic characteristics of beech (Fagus sylvatica) saplings during three years of exposure to elevated CO(2) concentration

Beech (Fagus sylvatica L.) seedlings were cultivated from seeds sown in pots or directly in the ground in outdoor chambers that were transparent to solar radiation, and provided either ambient air or CO(2)-enriched air (ambient + 350 &mgr;mol mol(-1)). The rooting volume was high in all experiments. In the short-term experiment, potted plants were assigned to a factorial CO(2) x nutrient treatment (optimal nutrient supply and severe nutrient shortage) for 1 year. In the long-term experiment, plants were grown directly in the ground and received an optimal supply of water and nutrients in both CO(2) treatments for 3 years. Nutrient stress caused carboxylation capacity (V(m)) to decrease in the potted seedlings exposed to CO(2)-enriched air during their first growing season. In the long-term experiment with optimal nutrient supply, CO(2)-enriched air did not affect V(m), but caused an upward acclimation of maximum electron transport rate (J(m)). Consequently, there was a 14% increase in the J(m)/V(m) ratio, indicating nitrogen reallocation to maintain an equilibrium between RuBP consumption and RuBP regeneration. Both V(m) and J(m) decreased during the growing season in both CO(2) treatments. Although upward acclimation of J(m) was no longer apparent at the end of the third growing season, plants in CO(2)-enriched air maintained a higher J(m)/V(m) ratio than plants in ambient air, indicating that photosynthetic acclimation always occurred. Second flush leaves appeared during each growing season. When expressed on the basis of foliar nitrogen concentration, their photosynthetic characteristics (V(m) and J(m)) were enhanced compared with other leaves. Because the number of second flush leaves was also increased in the elevated CO(2) treatment, this response should be taken into account when modeling the effects of elevated CO(2) concentration on canopy photosynthesis. Stomatal conductance decreased in response to atmospheric CO(2) enrichment; however, the stomatal response to irradiance followed a single relationship based on two stomatal conductance models.     

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.     

Photosynthetic responses to needle water potentials in Scots pine after a four-year exposure to elevated CO(2) and temperature

Effects of needle water potential (Psi(l)) on gas exchange of Scots pine (Pinus sylvestris L.) grown for 4 years in open-top chambers with elevated temperature (ET), elevated CO(2) (EC) or a combination of elevated temperature and CO(2) (EC + ET) were examined at a high photon flux density (PPFD), saturated leaf to air water vapor pressure deficit (VPD) and optimal temperature (T). We used the Farquhar model of photosynthesis to estimate the separate effects of Psi(l) and the treatments on maximum carboxylation efficiency (V(c,max)), ribulose-1,5-bisphosphate regeneration capacity (J), rate of respiration in the light (R(d)), intercellular partial pressure of CO(2) (C(i)) and stomatal conductance (G(s)). Depression of CO(2) assimilation rate at low Psi(l) was the result of both stomatal and non-stomatal limitations on photosynthetic processes; however, stomatal limitations dominated during short-term water stress (Psi(l) < -1.2 MPa), whereas non-stomatal limitations dominated during severe water stress. Among the nonstomatal components, the decrease in J contributed more to the decline in photosynthesis than the decrease in V(c,max). Long-term elevation of CO(2) and temperature led to differences in the maximum values of the parameters, the threshold values of Psi(l) and the sensitivity of the parameters to decreasing Psi(l). The CO(2) treatment decreased the maximum values of V(c,max), J and R(d) but significantly increased the sensitivity of V(c,max), J and R(d) to decreasing Psi(l) (P < 0.05). The effects of the ET and EC + ET treatments on V(c,max), J and R(d) were opposite to the effects of the EC treatment on these parameters. The values of G(s), which were measured simultaneously with maximum net rate of assimilation (A(max)), declined in a curvilinear fashion as Psi(l) decreased. Both the EC + ET and ET treatments significantly decreased the sensitivity of G(s) to decreasing Psi(l). We conclude that, in the future, acclimation to increased atmospheric CO(2) and temperature could increase the tolerance of Scots pine to water stress.     

Seedling response of two trembling aspen (Populus tremuloides) families to infection by Hypoxylon mammatum

Open-pollinated progenies from two trembling aspen (Populus tremuloides) families with a 30-year field history of 91% (putatively susceptible, PS) and 45% (putatively resistant, PR) mortality, respectively, caused by Hypoxylon mammatum, were grown in a greenhouse for 3 and 9 months. Seedlings were inoculated with two isolates of H. mammatum, representing high and low aggressiveness, through a stem wound. Beginning 4 days after inoculation, host responses were examined on 312 seedlings per family for a period of 5 weeks on 3- and 9-month-old seedlings. Significant differences in host responses were observed between the two isolates, with the more aggressive isolate allowing less callus formation, causing more tissue necrosis and producing longer cankers than the less aggressive isolate on both aspen families. Between-family difference was much greater than within-family difference for tissue necrosis, callus formation and canker length for both seedling ages. When the 3-month-old seedlings were challenged with H. mammatum, the PR family showed greater tissue necrosis, longer cankers, and less callus formation than the PS family. However, when the more lignifìed 9-month-old seedlings of the same families were inoculated, the PR seedlings had much less tissue necrosis, shorter cankers, more callus formation and less seedling mortality than the PS family. The results indicate that artificial inoculation of seedlings in a greenhouse can differentiate two aspen families at the seedling stage, but their correlation to field performance is highly dependent upon the seedling age. Host responses of older seedlings may be a better indicator of field performance.     

Comparison of the photosynthetic capacity of Siberian and Japanese birch seedlings grown in elevated CO(2) and temperature

The effects of increased CO(2) and temperature on the photosynthetic capacity of Siberian white birch and Japanese white birch (Betula platyphylla Sukatch. and B. platyphylla Sukatch. var. japonica Hara) were measured. Birch seedlings were raised with a CO(2) partial pressure of 36 +/- 0.3 Pa (i.e., ambient) or 70 +/- 0.6 Pa at day/night temperatures of either 30/16 degrees C or 26/12 degrees C. Siberian birch leaves were smaller and thicker than Japanese birch leaves. Water use efficiency and nitrogen use efficiency of Siberian birch grown in the CO(2)-enriched air were higher than those of Japanese birch. Both species showed a physiological adjustment to the growth CO(2) partial pressure. Carboxylation efficiency and quantum yield of both species grown in CO(2)-enriched air were lower than those of seedlings grown in ambient CO(2). The adaptation of Siberian and Japanese birch to elevated CO(2) and temperature are discussed in relation to predicted climate change.     

Responses of foliar gas exchange to long-term elevated CO(2) concentrations in mature loblolly pine trees

Branches of field-grown mature loblolly pine (Pinus taeda L.) trees were exposed for 2 years (1992 and 1993) to ambient or elevated CO(2) concentrations (ambient + 165 micro mol mol(-1) or ambient + 330 micro mol mol(-1) CO(2)). Exposure to elevated CO(2) concentrations enhanced rates of net photosynthesis (P(n)) by 53-111% compared to P(n) of foliage exposed to ambient CO(2). At the same CO(2) measurement concentration, the ratio of intercellular to atmospheric CO(2) concentration (C(i)/C(a)) and stomatal conductance to water vapor did not differ among foliage grown in an ambient or enriched CO(2) concentration. Analysis of the relationship between P(n) and C(i) indicated no significant change in carboxylation efficiency of ribulose-1,5-bisphosphate carboxylase/oxygenase during growth in elevated CO(2) concentrations. Based on estimates derived from P(n)/C(i) curves, there were no apparent treatment differences in dark respiration, CO(2) compensation point or P(n) at the mean C(i). In 1992, foliage in the three CO(2) treatments yielded similar estimates of CO(2)-saturated P(n) (P(max)), whereas in 1993, estimates of P(max) were higher for branches grown in elevated CO(2) than in ambient CO(2). We conclude that field-grown loblolly pine trees do not exhibit downward acclimation of leaf-level photosynthesis in their long-term response to elevated CO(2) concentrations.     

Atmospheric CO(2) and the ratio of intercellular to ambient CO(2) concentrations in plants

Much attention is focused today on predicting how plants will respond to anticipated changes in atmospheric composition and climate, and in particular to increases in CO(2) concentration. Here we review the long-term global fluctuations in atmospheric CO(2) concentration as a framework for understanding how current trends in atmospheric CO(2) concentration fit into a selective, evolutionary context. We then focus on an integrated approach for understanding how gas exchange metabolism responds to current environmental conditions, how it previously responded to glacial-interglacial conditions, and how it may respond to future changes in atmospheric CO(2) concentration.     

Ecophysiological responses of woody plants to past CO(2) concentrations

An approach is detailed for calculating historical rates of CO(2) uptake and water loss of leaves from measurements of leaf delta(13)C composition and climatic information. This approach was applied to investigate leaf gas exchange metabolism of woody taxa during the past 200 years of atmospheric CO(2) increase and in response to the longer-term atmospheric CO(2) increases plants experienced over the Pleistocene. Reconstructed net assimilation rates and water use efficiencies increased in response to increasing atmospheric CO(2) concentrations in both sets of material, whereas stomatal conductance, showing the combined responses of changes in stomatal density and leaf assimilation rates, was generally less responsive. Woody temperate taxa maintained a nearly constant c(i)/c(a) ratio in response to the increase in atmospheric CO(2) concentrations over both timescales, in part, as a result of changes in stomatal density. The reconstructed leaf-scale physiological responses to past global climatic and atmospheric change corroborated those anticipated from experimental work indicating the adequate capacity of experiments, at least at the scale of individual leaves, to predict plant responses to future environmental change.     

Long-term photosynthetic acclimation to increased atmospheric CO(2) concentration in young birch (Betula pendula) trees

To study the long-term response of photosynthesis to elevated atmospheric CO(2) concentration in silver birch (Betula pendula Roth.), 18 trees were grown in the field in open-top chambers supplied with 350 or 700 &mgr;mol mol(-1) CO(2) for four consecutive growing seasons. Maximum photosynthetic rates, stomatal conductance and CO(2) response curves were measured over the fourth growing season with a portable photosynthesis system. The photosynthesis model developed by Farquhar et al. (1980) was fitted to the CO(2) response curves. Chlorophyll, soluble proteins, total nonstructural carbohydrates, nitrogen and Rubisco activity were determined monthly. Elevated CO(2) concentration stimulated photosynthesis by 33% on average over the fourth growing season. However, comparison of maximum photosynthetic rates at the same CO(2) concentration (350 or 700 &mgr;mol mol(-1)) revealed that the photosynthetic capacity of trees grown in an elevated CO(2) concentration was reduced. Analysis of the response curves showed that acclimation to elevated CO(2) concentration involved decreases in carboxylation efficiency and RuBP regeneration capacity. No clear evidence for a redistribution of nitrogen within the leaf was observed. Down-regulation of photosynthesis increased as the growing season progressed and appeared to be related to the source-sink balance of the trees. Analysis of the main leaf components revealed that the reduction in photosynthetic capacity was accompanied by an accumulation of starch in leaves (100%), which was probably responsible for the reduction in Rubisco activity (27%) and to a lesser extent for reductions in other photosynthetic components: chlorophyll (10%), soluble protein (9%), and N concentrations (12%) expressed on an area basis. Despite a 21% reduction in stomatal conductance in response to the elevated CO(2) treatment, stomatal limitation was significantly less in the elevated, than in the ambient, CO(2) treatment. Thus, after four growing seasons exposed to an elevated CO(2) concentration in the field, the trees maintained increased photosynthetic rates, although their photosynthetic capacity was reduced compared with trees grown in ambient CO(2).     

Responses of loblolly pine seedlings to elevated CO(2) and fluctuating water supply

Osmotic adjustment of loblolly pine (Pinus taeda L.) seedlings to fluctuating water supply in elevated CO(2) was investigated. Seedlings were grown in controlled-environment chambers in either 350 or 700 micro l l(-1) CO(2) with weekly watering for four months, after which they were either watered weekly (well-watered treatment) or every two weeks (water-stress treatment) for 59 days. Osmotic adjustment was assessed by pressure-volume analysis of shoots and by analysis of soluble carbohydrates and free amino acids in roots during the last drying cycle. In well-watered seedlings, elevated CO(2) increased the concentration of soluble sugars in roots by 68%. Water stress reduced the soluble sugar concentration in roots of seedling growing in ambient CO(2) to 26% of that in roots of well-watered seedlings. Elevated CO(2) mitigated the water stress-induced decrease in the concentration of soluble sugars in roots. However, this was probably due, in part, to carbohydrate loading during the first four months when all seedlings were grown in the presence of a high water supply, rather than to osmotic adjustment to water stress. Water stress caused a doubling in the concentration of free primary amino acids in roots, whereas elevated CO(2) reduced primary amino acid and nitrogen concentrations to 32 and 74%, respectively, of those in roots of seedlings grown in ambient CO(2). There was no indication of large-scale osmotic adjustment to water stress or that elevated CO(2) enhanced osmotic adjustment in loblolly pine.     

Short-term effects of fertilization on photosynthesis and leaf morphology of field-grown loblolly pine following long-term exposure to elevated CO(2) concentration

We examined effects of a first nitrogen (N) fertilizer application on upper-canopy needle morphology and gas exchange in approximately 20-m-tall loblolly pine (Pinus taeda L.) exposed to elevated carbon dioxide concentration ([CO(2)]) for 9 years. Duke Forest free-air CO(2) enrichment (FACE) plots were split and half of each ring fertilized with 112 kg ha(-1) elemental N applied in two applications in March and April 2005. Measurements of needle length (L), mass per unit area (LMA), N concentration (N(l)) on a mass and an area basis, light-saturated net photosynthesis per unit leaf area (A(a)) and per unit mass (A(m)), and leaf conductance (g(L)) began after the second fertilizer application in existing 1-year-old foliage (F(O)) and later in developing current-year first-flush (F(C1)) and current-year second-flush (F(C2)) foliage. Elevated [CO(2)] increased A(a) by 43 and 52% in F(O) and F(C1) foliage, respectively, but generally had no significant effect on any other parameter. Fertilization had little or no significant effect on L, LMA, A or g(L) in F(O) foliage; although N(l) was significantly higher in fertilized trees by midsummer. In contrast, fertilization resulted in large increases in L, N(l), and A in F(C1) and F(C2) foliage, increasing A(a) by about 20%. These results suggest that, although both needle age classes accumulate N following fertilization, they use it differently-current-year foliage incorporates N into photosynthetic machinery, whereas 1-year-old foliage serves as an N store. There were no significant interaction effects of elevated [CO(2)] and fertilization on A. Elevated [CO(2)] increased the intercept of the A:N(l) relationship but did not significantly affect the slope of the relationship in either foliage age class.     

Sap flux in pure aspen and mixed aspen-birch forests exposed to elevated concentrations of carbon dioxide and ozone

Elevated concentrations of atmospheric carbon dioxide ([CO2]) and tropospheric ozone ([O3]) have the potential to affect tree physiology and structure and hence forest water use, which has implications for climate feedbacks. We investigated how a 40% increase above ambient values in [CO2] and [O3], alone and in combination, affect tree water use of pure aspen and mixed aspen-birch forests in the free air CO2-O3 enrichment experiment near Rhinelander, Wisconsin (Aspen FACE). Measurements of sap flux and canopy leaf area index (L) were made during two growing seasons, when steady-state L had been reached after more than 6 years of exposure to elevated [CO2] and [O3]. Maximum stand-level sap flux was not significantly affected by elevated [O3], but was increased by 18% by elevated [CO2] averaged across years, communities and O(3) regimes. Treatment effects were similar in pure aspen and mixed aspen-birch communities. Increased tree water use in response to elevated [CO2] was related to positive CO2 treatment effects on tree size and L (+40%). Tree water use was not reduced by elevated [O3] despite strong negative O3 treatment effects on tree size and L (-22%). Elevated [O3] predisposed pure aspen stands to drought-induced sap flux reductions, whereas increased tree water use in response to elevated [CO2] did not result in lower soil water content in the upper soil or decreasing sap flux relative to control values during dry periods. Maintenance of soil water content in the upper soil in the elevated [CO2] treatment was at least partly a function of enhanced soil water-holding capacity, probably a result of increased organic matter content from increased litter inputs. Our findings that larger trees growing in elevated [CO2] used more water and that tree size, but not maximal water use, was negatively affected by elevated [O3] suggest that the long-term cumulative effects on stand structure may be more important than the expected primary stomatal closure responses to elevated [CO2] and [O3] in determining stand-level water use under possible future atmospheric conditions.