Advances in understanding ozone impact on forest trees: messages from novel phytotron and free-air fumigation studies.

Recent evidence from novel phytotron and free-air ozone (O3) fumigation experiments in Europe and America on forest tree species is highlighted in relation to previous chamber studies. Differences in O3 sensitivity between pioneer and climax species are examined and viewed for trees growing at the harsh alpine timberline ecotone. As O3 apparently counteracts positive effects of elevated CO2 and mitigates productivity increases, response is governed by genotype, competitors, and ontogeny rather than species per se. Complexity in O3 responsiveness increased under the influence of pathogens and herbivores. The new evidence does not conflict in principle with previous findings that, however, pointed to a low ecological significance. This new knowledge on trees' O3 responsiveness beyond the juvenile stage in plantations and forests nevertheless implies limited predictability due to complexity in biotic and abiotic interactions. Unravelling underlying mechanisms is mandatory for assessing O3 risks as an important component of climate change scenarios.

Effects of elevated atmospheric CO2 and tropospheric O3 on tree branch growth and implications for hydrologic budgeting.

The forest hydrologic budget may be impacted by increasing CO(2) and tropospheric O(3). Efficient means to quantify such effects are beneficial. We hypothesized that changes in the balance of canopy interception, stem flow, and through-fall in the presence of elevated CO(2) and O(3) could be discerned using image analysis of leafless branches. We compared annual stem flow to the results of a computerized analysis of all branches from the 2002, 2004, and 2006 annual growth whorls of 97 ten-year-old trees from the Aspen Free-Air CO(2) and O(3) Enrichment (Aspen FACE) experiment in Rhinelander, WI. We found significant effects of elevated CO(2) and O(3) on some branch metrics, and that the branch metrics were useful for predicting stem flow from birch, but not aspen. The results of this study should contribute to development of techniques for efficient characterization of effects on the forest hydrologic budget of increasing CO(2) and tropospheric O(3).

Effects of elevated atmospheric CO2 and/or O3 on intra- and interspecific competitive ability of aspen.

Three model communities of trembling aspen (monoculture, and mixed with either paper birch or sugar maple) were grown for seven years in elevated atmospheric CO(2) and O(3) using Free Air CO(2) Enrichment (FACE) technology. We utilized trends in species' importance, calculated as an index of volume growth and survival, as indications of shifting community composition. For the pure aspen communities, different clones emerged as having the highest change in relative importance values depending on the pollutant exposure. In the control and elevated CO(2) treatments, clone 42E was rapidly becoming the most successful clone while under elevated O(3), clone 8 L emerged as the dominant clone. In fact, growth of clone 8 L was greater in the elevated O(3) treatment compared to controls. For the mixed aspen-birch community, importance of aspen and birch changed by - 16 % and + 62 %, respectively, in the controls. In the treatments, however, importance of aspen and birch changed by - 27 % and + 87 %, respectively, in elevated O(3), and by - 10 % and + 45 %, respectively, in elevated CO(2). Thus, the presence of elevated O(3) hastened conversion of stands to paper birch, whereas the presence of elevated CO(2) delayed it. Relative importance of aspen and maple changed by - 2 % and + 3 %, respectively, after seven years in the control treatments. But in elevated O(3), relative importance of aspen and maple changed by - 2 % and + 5 %, respectively, and in elevated CO(2) by + 9 and - 20 %, respectively. Thus, elevated O(3) slightly increases the rate of conversion of aspen stands to sugar maple, but maple is placed at a competitive disadvantage to aspen under elevated CO(2).

Combined effects of atmospheric CO2 and N availability on the belowground carbon and nitrogen dynamics of aspen mesocosms.

It is uncertain whether elevated atmospheric CO2 will increase C storage in terrestrial ecosystems without concomitant increases in plant access to N. Elevated CO2 may alter microbial activities that regulate soil N availability by changing the amount or composition of organic substrates produced by roots. Our objective was to determine the potential for elevated CO2 to change N availability in an experimental plant-soil system by affecting the acquisition of root-derived C by soil microbes. We grew Populus tremuloides (trembling aspen) cuttings for 2 years under two levels of atmospheric CO2 (36.7 and 71.5 Pa) and at two levels of soil N (210 and 970 µg N g-1). Ambient and twice-ambient CO2 concentrations were applied using open-top chambers, and soil N availability was manipulated by mixing soils differing in organic N content. From June to October of the second growing season, we measured midday rates of soil respiration. In August, we pulse-labeled plants with 14CO2 and measured soil 14CO2 respiration and the 14C contents of plants, soils, and microorganisms after a 6-day chase period. In conjunction with the August radio-labeling and again in October, we used 15N pool dilution techniques to measure in situ rates of gross N mineralization, N immobilization by microbes, and plant N uptake. At both levels of soil N availability, elevated CO2 significantly increased whole-plant and root biomass, and marginally increased whole-plant N capital. Significant increases in soil respiration were closely linked to increases in root biomass under elevated CO2. CO2 enrichment had no significant effect on the allometric distribution of biomass or 14C among plant components, total 14C allocation belowground, or cumulative (6-day) 14CO2 soil respiration. Elevated CO2 significantly increased microbial 14C contents, indicating greater availability of microbial substrates derived from roots. The near doubling of microbial 14C contents at elevated CO2 was a relatively small quantitative change in the belowground C cycle of our experimental system, but represents an ecologically significant effect on the dynamics of microbial growth. Rates of plant N uptake during both 6-day periods in August and October were significantly greater at elevated CO2, and were closely related to fine-root biomass. Gross N mineralization was not affected by elevated CO2. Despite significantly greater rates of N immobilization under elevated CO2, standing pools of microbial N were not affected by elevated CO2, suggesting that N was cycling through microbes more rapidly. Our results contained elements of both positive and negative feedback hypotheses, and may be most relevant to young, aggrading ecosystems, where soil resources are not yet fully exploited by plant roots. If the turnover of microbial N increases, higher rates of N immobilization may not decrease N availability to plants under elevated CO2.

Populus tremuloides photosynthesis and crown architecture in response to elevated CO2 and soil N availability.

We tested the hypothesis that elevated CO2 would stimulate proportionally higher photosynthesis in the lower crown of Populus trees due to less N retranslocation, compared to tree crowns in ambient CO2. Such a response could increase belowground C allocation, particularly in trees with an indeterminate growth pattern such as Populus tremuloides. Rooted cuttings of P. tremuloides were grown in ambient and twice ambient (elevated) CO2 and in low and high soil N availability (89 ± 7 and 333 ± 16 ng N g-1 day-1 net mineralization, respectively) for 95 days using open-top chambers and open-bottom root boxes. Elevated CO2 resulted in significantly higher maximum leaf photosynthesis (A max) at both soil N levels. A max was higher at high N than at low N soil in elevated, but not ambient CO2. Photosynthetic N use efficiency was higher at elevated than ambient CO2 in both soil types. Elevated CO2 resulted in proportionally higher whole leaf A in the lower three-quarters to one-half of the crown for both soil types. At elevated CO2 and high N availability, lower crown leaves had significantly lower ratios of carboxylation capacity to electron transport capacity (V cmax/J max) than at ambient CO2 and/or low N availability. From the top to the bottom of the tree crowns, V cmax/J max increased in ambient CO2, but it decreased in elevated CO2 indicating a greater relative investment of N into light harvesting for the lower crown. Only the mid-crown leaves at both N levels exhibited photosynthetic down regulation to elevated CO2. Stem biomass segments (consisting of three nodes and internodes) were compared to the total A leaf for each segment. This analysis indicated that increased A leaf at elevated CO2 did not result in a proportional increase in local stem segment mass, suggesting that C allocation to sinks other than the local stem segment increased disproportionally. Since C allocated to roots in young Populus trees is primarily assimilated by leaves in the lower crown, the results of this study suggest a mechanism by which C allocation to roots in young trees may increase in elevated CO2.

Effects of elevated CO(2) and light availability on the photosynthetic light response of trees of contrasting shade tolerance.

Photosynthetic light response curves (A/PPFD), leaf N concentration and content, and relative leaf absorbance (alpha(r)) were measured in 1-year-old seedlings of shade-intolerant Betula papyrifera Marsh., moderately shade-tolerant Quercus rubra L. and shade-tolerant Acer rubrum L. Seedlings were grown in full sun or 26% of full sun (shade) and in ambient (350 ppm) or elevated (714 ppm) CO(2) for 80 days. In the shade treatments, 80% of the daily PPFD on cloud-free days was provided by two 30-min sun patches at midday. In Q. rubra and A. rubrum, leaf N concentration and alpha(r) were significantly higher in seedlings in the shade treatments than in the sun treatments, and leaf N concentration was lower in seedlings in the ambient CO(2) treatments than in the elevated CO(2) treatments. Changes in alpha(r) and leaf N content suggest that reapportionment of leaf N into light harvesting machinery in response to shade and elevated CO(2) tended to increase with increasing shade tolerance of the plant. Shifts induced by elevated CO(2) in the A/PPFD relationship in sun plants were largest in B. papyrifera and least in A. rubrum: the reverse was true for shade plants. Elevated CO(2) resulted in increased light-saturated A in every species x light treatment combination, except in shaded B. papyrifera. The light compensation point (Gamma) decreased in response to shade in all species, and in response to elevated CO(2) in A. rubrum and Q. rubra. Acer rubrum had the greatest increases in apparent quantum yield (phi) in response to shade and elevated CO(2). To illustrate the effects of shifts in A, Gamma and phi on daily C gain, daily integrated C balance was calculated for individual sun and shade leaves. Ignoring possible stomatal effects, estimated daily (24 h) leaf C balance was 218 to 442% higher in the elevated CO(2) treatments than in the ambient CO(2) treatments in both sun and shade seedlings of Q. rubra and A. rubrum. These results suggest that the ability of species to acclimate photosynthetically to elevated CO(2) may, in part, be related to their ability to adapt to low irradiance. Such a relationship has implications for altered C balance and nitrogen use efficiency of understory seedlings.

Ecophysiological and morphological responses to shade and drought in two contrasting ecotypes of Prunus serotina.

Photosynthesis (A), water relations and stomatal reactivity during drought, and leaf morphology were evaluated on 2-year-old, sun- and shade-grown Prunus serotina Ehrh. seedlings of a mesic Pennsylvania seed source and a more xeric Wisconsin source. Wisconsin plants maintained higher A and leaf conductance (g(wv)) than Pennsylvania plants during the entire drought under sun conditions, and during the mid stages of drought under shade conditions. Compared to shade plants, sun plants of both sources exhibited a more rapid decrease in A or % A(max) with decreasing leaf water potential (Psi). Tissue water relations parameters were generally not significantly different between seed sources. However, osmotic potentials were lower in sun than shade plants under well-watered conditions. Following drought, shade plants, but not sun plants, exhibited significant osmotic adjustment. Sun leaves had greater thickness, specific mass, area and stomatal density and lower guard cell length than shade leaves in one or both sources. Wisconsin sun leaves were seemingly more xerophytic with greater thickness, specific mass, and guard cell length than Pennsylvania sun leaves. No source differences in leaf structure were exhibited in shade plants. Stomatal reactivity to sun-shade cycles was similar between ecotypes. However, well-watered and droughted plants differed in stomatal reactivity within and between multiple sun-shade cycles. The observed ecotypic and phenotypic variations in ecophysiology and morphology are consistent with the ability of Prunus serotina to survive in greatly contrasting environments.

Rehydration effects on pressure-volume relationships in four temperate woody species: variability with site, time of season and drought conditions.

Seasonal pressure-volume (P-V) analyses were conducted on rehydrated and non-rehydrated leaves of Quercus rubra, Q. ilicifolia, Q. prinus, and Fraxinus americana in central Pennsylvania, U.S.A., to test the hypothesis that rehydration-induced shifts in P-V parameters occur in woody species from a non-arid region, and that the magnitude of these shifts increases with species drought tolerance and drought conditions. The species from a xeric ridge (Q. ilicifolia and Q. prinus) displayed increases of about 0.4-0.6 MPa in the osmotic potentials at full and zero turgor and a concurrent loss of symplastic solutes following 12 h and 24 h rehydration, particularly during a late-season drought. In contrast, the mesic, valley species (Q. rubra and F. americana) did not display significant shifts in osmotic parameters with rehydration at any time. In several instances, the relative water content at zero turgor (RWC0) increased by about 6% (e.g., from 85% to 91%) and the bulk elastic modulus (ε) decreased by about 4.0 MPa following rehydration and correction for the plateau effect; the magnitude of these shifts was greatest in the xeric species. However, when data were not corrected for the plateau effect, RWC0 decreased by about 4% in some of the species/date combinations. Plateaus were also responsible for some of the decrease in ε with rehydration, but not for the shifts in osmotic potentials. The largest increases in osmotic potentials corresponded with decreases in tissue osmotic solute content. Rehydration-induced shifts in P-V parameters were responsible for masking or reducing most of the species and seasonal differences exhibited in nonrehydrated samples.

Drought adaptations and responses in five genotypes of Fraxinus pennsylvanica Marsh.: photosynthesis, water relations and leaf morphology.

Genotypic variation in photosynthesis and plant water relations during drought, and in leaf and seedling morphology were examined in greenhouse-grown Fraxinus pennsylvanica Marsh. (green ash) from five populations located along an east-west transect from New York State to South Dakota. During a 17-day drought, South Dakota seedlings, from the most xeric habitat, maintained the highest net photosynthesis and leaf conductance, and New York seedlings, from the most mesic habitat, exhibited the lowest net photosynthesis and leaf conductance. All populations except New York adjusted osmotically during the 17-day drought, by the end of which New York seedlings had the highest osmotic potentials at full and zero turgor. Tissue elasticity increased in New York seedlings, but decreased in Nebraska seedlings during the drought. Leaves of South Dakota seedlings were the most xerophytic. They were smaller in area and greater in thickness and specific mass than leaves of other sources. Leaves of New York seedlings were thinner than those of the other genotypes and among the largest. Seedlings from South Dakota were smaller than those of the other populations.