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Three young northern temperate forest communities in the north-central United States were exposed to factorial combinations of elevated carbon dioxide ( CO2) and tropospheric ozone (O3) for 11 years. Here, we report results from an extensive sampling of plant biomass and soil conducted at the conclusion of the experiment that enabled us to estimate ecosystem carbon (C) content and cumulative net primary productivity ( NPP). Elevated CO2 enhanced ecosystem C content by 11%, whereas elevated O3 decreased ecosystem C content by 9%. There was little variation in treatment effects on C content across communities and no meaningful interactions between CO2 and O3. Treatment effects on ecosystem C content resulted primarily from changes in the near-surface mineral soil and tree C, particularly differences in woody tissues. Excluding the mineral soil, cumulative NPP was a strong predictor of ecosystem C content ( r 2 = 0.96). Elevated CO2 enhanced cumulative NPP by 39%, a consequence of a 28% increase in canopy nitrogen (N) content (g N m−2) and a 28% increase in N productivity ( NPP/canopy N). In contrast, elevated O3 lowered NPP by 10% because of a 21% decrease in canopy N, but did not impact N productivity. Consequently, as the marginal impact of canopy N on NPP (∆ NPP/∆N) decreased through time with further canopy development, the O3 effect on NPP dissipated. Within the mineral soil, there was less C in the top 0.1 m of soil under elevated O3 and less soil C from 0.1 to 0.2 m in depth under elevated CO2. Overall, these results suggest that elevated CO2 may create a sustained increase in NPP, whereas the long-term effect of elevated O3 on NPP will be smaller than expected. However, changes in soil C are not well-understood and limit our ability to predict changes in ecosystem C content. [ABSTRACT FROM AUTHOR]

Summary 1. Competition effects on growth of individual trees were examined for 4 years in aggrading, mixed-clone stands of trembling aspen (Populus tremuloides Michx.) at the Aspen-FACE free-air CO2 and O3 enrichment facility in northern Wisconsin, USA. During each growing season stands received one of four combinations of atmospheric [CO2 ] (ambient vs ∼56 Pa) and [O3 ] (ambient vs ∼1·5 × ambient). 2. Non-destructive measurements of annual tree growth were compared within and among clones and treatments in relation to an index of competitive status based on the difference between a tree's height and that of its four nearest neighbours. Competitive status strongly influenced tree growth, and the positive growth response to elevated [CO2 ] was greater for competitively advantaged individuals than for disadvantaged individuals of most clones. 3. The magnitude of O3 effects on growth depended on clone and competitive status: for some clones, negative O3 effects were stronger with competitive advantage while others showed stronger O3 effects with competitive disadvantage. The interactive effects of CO2 and O3 differed among clones, with negative effects of O3 amplified or ameliorated by elevated CO2 , depending on clone and competitive status. 4. Treatments modified competitive interactions by affecting the magnitude of growth differences among clones. These modifications did not alter clone rankings of competitive performance, but when CO2 and O3 were both elevated, the differences in competitive performance among clones decreased. [ABSTRACT FROM AUTHOR]

Abstract Leaf photosynthesis (Ps), nitrogen (N) and light environment were measured on Populus tremuloides trees in a developing canopy under free-air CO2 enrichment in Wisconsin, USA. After 2 years of growth, the trees averaged 1·5 and 1·6 m tall under ambient and elevated CO2, respectively, at the beginning of the study period in 1999. They grew to 2·6 and 2·9 m, respectively, by the end of the 1999 growing season. Daily integrated photon flux from cloud-free days (PPFDday,sat) around the lowermost branches was 16·8 ± 0·8 and 8·7 ± 0·2% of values at the top for the ambient and elevated CO2 canopies, respectively. Elevated CO2 significantly decreased leaf N on a mass, but not on an area, basis. N per unit leaf area was related linearly to PPFDday,sat throughout the canopies, and elevated CO2 did not affect that relationship. Leaf Ps light-response curves responded differently to elevated CO2, depending upon canopy position. Elevated CO2 increased Pssat only in the upper (unshaded) canopy, whereas characteristics that would favour photosynthesis in shade were unaffected by elevated CO2. Consequently, estimated daily integrated Ps on cloud-free days (Psday,sat) was stimulated by elevated CO2 only in the upper canopy. Psday,sat of the lowermost branches was actually lower with elevated CO2 because of the darker light environment. The lack of CO2 stimulation at the mid- and lower canopy was probably related to significant down-regulation of photosynthetic capacity; there was no down-regulation of Ps in the upper canopy. The relationship between Psday,sat and leaf N indicated that N was not optimally allocated within the canopy in a manner that would maximize whole-canopy Ps or photosynthetic N use efficiency. Elevated CO2 had no effect on the optimization of canopy N allocation. [ABSTRACT FROM AUTHOR]

ABSTRACTLeaf gas exchange parameters and the content of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) in the leaves of two 2-year-old aspen (Populus tremuloides Michx.) clones (no. 216, ozone tolerant and no. 259, ozone sensitive) were determined to estimate the relative stomatal and mesophyll limitations to photosynthesis and to determine how these limitations were altered by exposure to elevated CO2 and/or O3. The plants were exposed either to ambient air (control), elevated CO2 (560 p.p.m.) elevated O3 (55 p.p.b.) or a mixture of elevated CO2 and O3 in a free air CO2 enrichment (FACE) facility located near Rhinelander, Wisconsin, USA. Light-saturated photosynthesis and stomatal conductance were measured in all leaves of the current terminal and of two lateral branches (one from the upper and one from the lower canopy) to detect possible age-related variation in relative stomatal limitation (leaf age is described as a function of leaf plastochron index). Photosynthesis was increased by elevated CO2 and decreased by O3 at both control and elevated CO2. The relative stomatal limitation to photosynthesis (ls) was in both clones about 10% under control and elevated O3. Exposure to elevated CO2 + O3 in both clones and to elevated CO2 in clone 259, decreased ls even further – to about 5%. The corresponding changes in Rubisco content and the stability of Ci/Ca ratio suggest that the changes in photosynthesis in response to elevated CO2 and O3 were primarily triggered by altered mesophyll processes in the two aspen clones of contrasting O3 tolerance. The changes in stomatal conductance seem to be a secondary response, maintaining stable Ci under the given treatment, that indicates close coupling between stomatal and mesophyll processes. [ABSTRACT FROM AUTHOR]

Seasonal patterns of [14C]‐labeled photosynthate distribution within two intensively cultured Populus clones with contrasting phenology (P. tristis × P. balsamifera cv. ‘Tristis no. 1′; P. × euramericana cv. Eugenei) were investigated during the establishment year. During active shoot elongation upper mature leaves exported 14C acropetally to the expanding leaves and elongating internodes, and basipetally to the stem. Little 14C was exported to lower mature leaves or lateral branches. At budset the 14C export pattern shifted dramatically in the basipetal direction, i.e., to the lower stem, hardwood cutting, and roots. The timing of budset was the primary factor determining the differences between the clones, except that in all cases Tristis exported more 14C to the roots than Eugenei. After budset lower mature leaves had a similar export pattern to upper leaves, but the quantity of 14C exported to the roots was slightly higher. The results confirm the importance of autumn foliage for root growth in poplar. Clonal differences in seasonal patterns of photosynthate distribution offer potential for the poplar breeder seeking to match a clone's growth pattern with the specific growing season of the site. [ABSTRACT FROM AUTHOR]

Variable internal plant nutrient content may confound plant response to environmental stress. Plant nutrient content may be controlled with relative addition rate techniques in solution culture. However, because raising large numbers of plants in flowing solution culture is difficult, we investigated the feasibility of raising plants in soil mix using relative fertilizer additions. Aspen (Populus tremuloides Michx.) clones (216, 259 and 271) planted in pots containing a peat, sand and vermiculite (2:1:1, v/v/v) soil mix were grown with exponentially increasing fertilizer concentrations and harvested periodically to assess growth. Addition rate treatments ranged from 0.01 to 0.05 day−1. The lag phase of growth, in which plants adjusted to the fertilizer regime, lasted 40 days after which plants entered the experimental period characterized by constant relative growth rates equivalent to applied fertilizer addition rates. Total plant nutrient concentration was (1) unique for each addition rate, (2) linearly related to addition rate and growth rate, and (3) tended to increase at the highest, and decrease at the lowest addition rates. Regardless, the plants appeared to have attained steady‐state conditions. Allocation of carbon to roots increased with lower addition rate treatments and was not dependent upon ontogeny. There were no treatment differences in growth response among aspen clones. Yet there were treatment differences in leaf chlorophyll and photosynthesis within the clones. For the 0.05 day−1 addition rate treatment, chlorophyll, leaf N concentration and photosynthetic rate were strongly correlated with one another, were at a maximum in recently mature leaves, and rapidly declined with leaf age. The rate of decline in these leaf characteristics was slowest in clone 271, consistent with the leaf longevity stress response reported elsewhere. Plant responses from these relative fertilizer addition trials in soil mix agree closely with those run in hydroponics, indicating that steady‐state nutrition can be achieved with a technically simple experimental assemblage. [ABSTRACT FROM AUTHOR]

Ethylene losses from the gas phase of various container-seal systems were studied to develop acceptable methods for containing ethylene during experiments. Ethylene at an initial amount of 104 μl l-1 was stored in glass vials at near atmospheric pressure for 20 h at 25-27°C and at 35% relative humidity external to the vials. Crimped serum vials sealed with saturated (NH4)2SO4 solution, neoprene rubber septa, nitrile rubber (Hycar) septa, butyl rubber septa, and brown translucent silicone rubber septa lost ethylene at the rate of 1.8. 10.2, 16.2, 16.5, and 40.2 nl m-2s-1, respectively, over the 20-h period. Screw-capped reaction vials sealed with white silicone rubber septa lost ethylene at the rate of 30.2 nl -2s-1. The (NH4)2SO4 solution was utilized as a seal by inverting a vial so that the salt solution covered the internal surface of the vial septum. Saturated (NH4)2SO4 solution is an effective seal. Silicone rubber should be avoided as a seal in systems for containing ethylene. Ethylene production values in the literature may be underestimates where silicone rubber seals have been used. [ABSTRACT FROM AUTHOR]

Abstract We investigated the interaction of elevated CO2 and/or (Ozone) O3 on the occurrence and severity of aspen leaf rust (Melampsora medusae Thuem. f. sp. tremuloidae ) on trembling aspen (Populus tremuloides Michx.). Furthermore, we examined the role of changes in leaf surface properties induced by elevated CO2 and/or O3 in this host–pathogen interaction. Three- to five-fold increases in levels of rust infection index were found in 2 consecutive years following growing-season-long exposures with either O3 alone or CO2 + O3 depending on aspen clone. Examination of leaf surface properties (wax appearance, wax amount, wax chemical composition, leaf surface and wettability) suggested significant effects by O3 and CO2 + O3 . We conclude that elevated O3 is altering aspen leaf surfaces in such a way that it is likely predisposing the plants to increased infection by aspen leaf rust. [ABSTRACT FROM AUTHOR]

ABSTRACTLeaf gas exchange parameters and the content of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) in the leaves of two 2-year-old aspen (Populus tremuloides Michx.) clones (no. 216, ozone tolerant and no. 259, ozone sensitive) were determined to estimate the relative stomatal and mesophyll limitations to photosynthesis and to determine how these limitations were altered by exposure to elevated CO2 and/or O3. The plants were exposed either to ambient air (control), elevated CO2 (560 p.p.m.) elevated O3 (55 p.p.b.) or a mixture of elevated CO2 and O3 in a free air CO2 enrichment (FACE) facility located near Rhinelander, Wisconsin, USA. Light-saturated photosynthesis and stomatal conductance were measured in all leaves of the current terminal and of two lateral branches (one from the upper and one from the lower canopy) to detect possible age-related variation in relative stomatal limitation (leaf age is described as a function of leaf plastochron index). Photosynthesis was increased by elevated CO2 and decreased by O3 at both control and elevated CO2. The relative stomatal limitation to photosynthesis (ls) was in both clones about 10% under control and elevated O3. Exposure to elevated CO2 + O3 in both clones and to elevated CO2 in clone 259, decreased ls even further – to about 5%. The corresponding changes in Rubisco content and the stability of Ci/Ca ratio suggest that the changes in photosynthesis in response to elevated CO2 and O3 were primarily triggered by altered mesophyll processes in the two aspen clones of contrasting O3 tolerance. The changes in stomatal conductance seem to be a secondary response, maintaining stable Ci under the given treatment, that indicates close coupling between stomatal and mesophyll processes. [ABSTRACT FROM AUTHOR]