Integrating ecophysiology and forest landscape models to improve projections of drought effects under climate change.

Fundamental drivers of ecosystem processes such as temperature and precipitation are rapidly changing and creating novel environmental conditions. Forest landscape models (FLM) are used by managers and policy-makers to make projections of future ecosystem dynamics under alternative management or policy options, but the links between the fundamental drivers and projected responses are weak and indirect, limiting their reliability for projecting the impacts of climate change. We developed and tested a relatively mechanistic method to simulate the effects of changing precipitation on species competition within the LANDIS-II FLM. Using data from a field precipitation manipulation experiment in a piñon pine (Pinus edulis) and juniper (Juniperus monosperma) ecosystem in New Mexico (USA), we calibrated our model to measurements from ambient control plots and tested predictions under the drought and irrigation treatments against empirical measurements. The model successfully predicted behavior of physiological variables under the treatments. Discrepancies between model output and empirical data occurred when the monthly time step of the model failed to capture the short-term dynamics of the ecosystem as recorded by instantaneous field measurements. We applied the model to heuristically assess the effect of alternative climate scenarios on the piñon-juniper ecosystem and found that warmer and drier climate reduced productivity and increased the risk of drought-induced mortality, especially for piñon. We concluded that the direct links between fundamental drivers and growth rates in our model hold great promise to improve our understanding of ecosystem processes under climate change and improve management decisions because of its greater reliance on first principles.

Wood properties of Populus and Betula in long-term exposure to elevated CO2 and O3.

We studied the interactive effects of elevated concentrations of CO2 and O3 on radial growth and wood properties of four trembling aspen (Populus tremuloides Michx.) clones and paper birch (Betula papyrifera Marsh.) saplings. The material for the study was collected from the Aspen FACE (free-air CO2 enrichment) experiment in Rhinelander (WI, USA). Trees had been exposed to four treatments [control, elevated CO2 (560 ppm), elevated O3 (1.5 times ambient) and combined CO2 + O3 ] during growing seasons 1998-2008. Most treatment responses were observed in the early phase of experiment. Our results show that the CO2- and O3-exposed aspen trees displayed a differential balance between efficiency and safety of water transport. Under elevated CO2, radial growth was enhanced and the trees had fewer but hydraulically more efficient larger diameter vessels. In contrast, elevated O3 decreased radial growth and the diameters of vessels and fibres. Clone-specific decrease in wood density and cell wall thickness was observed under elevated CO2 . In birch, the treatments had no major impacts on wood anatomy or wood density. Our study indicates that short-term impact studies conducted with young seedlings may not give a realistic view of long-term ecosystem responses.

Elevated carbon dioxide and ozone alter productivity and ecosystem carbon content in northern temperate forests.

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.

Can elevated CO2 and ozone shift the genetic composition of aspen (Populus tremuloides) stands?

The world's forests are currently exposed to increasing concentrations of carbon dioxide (CO2) and ozone (O3). Both pollutants can potentially exert a selective effect on plant populations. This, in turn, may lead to changes in ecosystem properties, such as carbon sequestration. Here, we report how elevated CO2 and O3 affect the genetic composition of a woody plant population via altered survival. Using data from the Aspen free-air CO2 enrichment (FACE) experiment (in which aspen clones were grown in factorial combinations of CO2 and O3), we develop a hierarchical Bayesian model of survival. We also examine how survival differences between clones could affect pollutant responses in the next generation. Our model predicts that the relative abundance of the tested clones, given equal initial abundance, would shift under either elevated CO2 or O3 as a result of changing survival rates. Survival was strongly affected by between-clone differences in growth responses. Selection could noticeably decrease O3 sensitivity in the next generation, depending on the heritability of growth responses and the distribution of seed production. The response to selection by CO2, however, is likely to be small. Our results suggest that the changing atmospheric composition could shift the genotypic composition and average pollutant responses of tree populations over moderate timescales.

Forest productivity under elevated CO2 and O3: positive feedbacks to soil N cycling sustain decade-long net primary productivity enhancement by CO2.

The accumulation of anthropogenic CO2 in the Earth's atmosphere, and hence the rate of climate warming, is sensitive to stimulation of plant growth by higher concentrations of atmospheric CO2. Here, we synthesise data from a field experiment in which three developing northern forest communities have been exposed to factorial combinations of elevated CO2 and O3. Enhanced net primary productivity (NPP) (c. 26% increase) under elevated CO2 was sustained by greater root exploration of soil for growth-limiting N, as well as more rapid rates of litter decomposition and microbial N release during decay. Despite initial declines in forest productivity under elevated O3, compensatory growth of O3 -tolerant individuals resulted in equivalent NPP under ambient and elevated O3. After a decade, NPP has remained enhanced under elevated CO2 and has recovered under elevated O3 by mechanisms that remain un-calibrated or not considered in coupled climate-biogeochemical models simulating interactions between the global C cycle and climate warming.

Acute O 3 damage on first year coppice sprouts of aspen and maple sprouts in an open-air experiment.

We studied the effect of high ozone (O(3)) concentration (110-490 nmol mol(-1)) on regenerating aspen (Populus tremuloides) and maple (Acer saccharum) trees at an open-air O(3) pollution experiment near Rhinelander WI USA. This study is the first of its kind to examine the effects of acute O(3) exposure on aspen and maple sprouts after the parent trees, which were grown under elevated O(3) and/or CO(2) for 12 years, were harvested. Acute O(3) damage was not uniform within the crowns of aspen suckers; it was most severe in the mature, fully expanded photosynthesizing leaves. Young expanding leaves showed no visible signs of acute O(3) damage contrary to expectations. Stomatal conductance played a primary role in the severity of acute O(3) damage as it directly controlled O(3) uptake. Maple sprouts, which had lower stomatal conductance, smaller stomatal aperture, higher stomatal density and larger leaf surface area, were tolerant of acute O(3) exposure. Moreover, elevated CO(2) did not ameliorate the adverse effects of acute O(3) dose on aspen and maple sprouts, in contrast to its ability to counteract the effects of long-term chronic exposure to lower O(3) levels.

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.

Wood properties of trembling aspen and paper birch after 5 years of exposure to elevated concentrations of CO(2) and O(3).

We investigated the interactive effects of elevated concentrations of carbon dioxide ([CO(2)]) and ozone ([O(3)]) on radial growth, wood chemistry and structure of five 5-year-old trembling aspen (Populus tremuloides Michx.) clones and the wood chemistry of paper birch (Betula papyrifera Marsh.). Material for the study was collected from the Aspen FACE (free-air CO(2) enrichment) experiment in Rhinelander, WI, where the saplings had been exposed to four treatments: control, elevated [CO(2)] (560 ppm), elevated [O(3)] (1.5 x ambient) and their combination for five growing seasons. Wood properties of both species were altered in response to exposure to the treatments. In aspen, elevated [CO(2)] decreased uronic acids (constituents of, e.g., hemicellulose) and tended to increase stem diameter. In response to elevated [O(3)] exposure, acid-soluble lignin concentration decreased and vessel lumen diameter tended to decrease. Elevated [O(3)] increased the concentration of acetone-soluble extractives in paper birch, but tended to decrease the concentration of these compounds in aspen. In paper birch, elevated [CO(2)] decreased and elevated [O(3)] increased starch concentration. The responses of wood properties to 5 years of fumigation differed from those previously reported after 3 years of fumigation.

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.

Impacts of elevated atmospheric CO2 and O3 on paper birch (Betula papyrifera): reproductive fitness.

Atmospheric CO2 and tropospheric O3 are rising in many regions of the world. Little is known about how these two commonly co-occurring gases will affect reproductive fitness of important forest tree species. Here, we report on the long-term effects of CO2 and O3 for paper birch seedlings exposed for nearly their entire life history at the Aspen FACE (Free Air Carbon Dioxide Enrichment) site in Rhinelander, WI. Elevated CO2 increased both male and female flower production, while elevated O3 increased female flower production compared to trees in control rings. Interestingly, very little flowering has yet occurred in combined treatment. Elevated CO2 had significant positive effect on birch catkin size, weight, and germination success rate (elevated CO2 increased germination rate of birch by 110% compared to ambient CO2 concentrations, decreased seedling mortality by 73%, increased seed weight by 17%, increased root length by 59%, and root-to-shoot ratio was significantly decreased, all at 3 weeks after germination), while the opposite was true of elevated O3 (elevated O3 decreased the germination rate of birch by 62%, decreased seed weight by 25%, and increased root length by 15%). Under elevated CO2, plant dry mass increased by 9 and 78% at the end of 3 and 14 weeks, respectively. Also, the root and shoot lengths, as well as the biomass of the seedlings, were increased for seeds produced under elevated CO2, while the reverse was true for seedlings from seeds produced under the elevated O3. Similar trends in treatment differences were observed in seed characteristics, germination, and seedling development for seeds collected in both 2004 and 2005. Our results suggest that elevated CO2 and O3 can dramatically affect flowering, seed production, and seed quality of paper birch, affecting reproductive fitness of this species.

Growth and C allocation of Populus tremuloides genotypes in response to atmospheric CO2 and soil N availability.

We grew cuttings of two early (mid Oct.) and two late (early Nov.) leaf-fall Populus tremuloides Michx. genotypes (referred to as genotype pairs) for c. 150 d in open-top chambers to understand how twice-ambient (elevated) CO2 and soil N availability would affect growth and C allocation. For the study, we selected genotypes differing in leaf area duration to find out if late-season photosynthesis influenced C allocation to roots. Both elevated CO2 and high soil N availability significantly increased estimated whole-tree photosynthesis, but they did so in different ways. Elevated CO2 stimulated leaf-level photosynthesis rates, whereas high soil N availability resulted in greater total plant leaf area. The early leaf-fall genotype pair had significantly higher photosynthesis rates per unit leaf area than the late leaf-fall genotype pair and elevated CO2 enhanced this difference. The early leaf-fall genotype pair had less leaf area than the late leaf-fall genotype pair, and their rate of leaf area development decreased earlier in the season. Across both genotype pairs, high soil N availability significantly increased fine root length production and mortality by increasing both the amount of root length present, and by decreasing the life span of individual roots. Elevated CO2 resulted in significantly increased fine root production and mortality in high N but not low N soil and did not affect fine root life span. The early leaf-fall genotype pair had significantly greater fine root length production than the late leaf-fall genotype pair across all CO2 and N treatments. These differences in belowground C allocations are consistent with the hypothesis that belowground C and N cycling is strongly influenced by soil N availability and will increase under elevated atmospheric CO2 . In addition, this study reinforces the need for better understanding of the variation in tree responses to elevated CO2 , within and among species.

Ecophysiological analysis of woody species in contrasting temperate communities during wet and dry years.

This study employed an intensive sampling regime in which leaf gas exchange and tissue-water relations were measured simultaneously on the same leaf at midday on 19 tree species from three distinct forest communities during wet (1990) and dry (1991) growing seasons. The study sites were located on a xeric barrens, a misic valley floor, and a wet-mesic floodplain in central Pennsylvania, United States. The xeric, mesic, and wetmesic sties had drought-related decreases in gravimetric soil moisture of 53, 34 and 27%, respectively. During the wet year, xeric and mesic communities had high seasonal mean photosynthetic rates (A) and stomatal conductance of water vapor (g wv) and low midday leaf water potential (ψ), whereas the wet-mesic community had low A and g wv and high midday ψ. The mesic and wet-mesic communities had dry year decreases in predawn ψ, g wv and A with the greatest drought effect occurring in the mesic community. Regression analysis indicated that species from each site that exhibited high wet-year A and g wv tended to have low midday ψ. This trend was reversed only in the mesic community in the drought year. Despite differences in midday ψ, all three communities had similar midday leaf turgor pressure (ψp) in the wet year attributable to lower osmotic potential at zero turgor (ψ π0 ) with increasing site droughtiness. Lower wet year ψ π0 in the xeric community was due to low symplast volume rather than high solute content. Species with the lowest ψ π0 in the wet year often did not have the lowest ψ π100 possibly related to differences in tissue elasticity. Moreover, increased elasticity during drought may have masked osmotic adjustment in ψ π100 but not in ψ π0 , via dilution of solutes at full hydration in some species. Despite the sampling regime used, there were no relationships between gas exchange and osmotic and elastic parameters that were consistently significant among communities or years. This result questions the universal, direct effect of osmotic and elastic adjustments in the maintenance of photosynthesis during drought. By including a large number of species, this study provided new insight to the ecophysiology of contrasting forest communities, and the community-wide impact of drought on contrasting sites.

Relationships among root branch order, carbon, and nitrogen in four temperate species.

The objective of this study was to examine how root length, diameter, specific root length, and root carbon and nitrogen concentrations were related to root branching patterns. The branching root systems of two temperate tree species, Acer saccharum Marsh. and Fraxinus americana L., and two perennial herbs from horizontal rhizomes, Hydrophyllum canadense L. and Viola pubescens Ait., were quantified by dissecting entire root systems collected from the understory of an A. saccharum-Fagus grandifolia Ehrh. forest. The root systems of each species grew according to a simple branching process, with laterals emerging from the main roots some distance behind the tip. Root systems normally consisted of only 4-6 branches (orders). Root diameter, length, and number of branches declined with increasing order and there were significant differences among species. Specific root length increased with order in all species. Nitrogen concentration increased with order in the trees, but remained constant in the perennial herbs. More than 75% of the cumulative root length of tree seedling root systems was accounted for by short (2-10 mm) lateral roots almost always <0.3 mm in diameter. Simple assumptions suggest that many tree roots normally considered part of the dynamic fine-root pool (e.g., all roots <2.0 mm in diameter) are too large to exhibit rapid rates of production and mortality. The smallest tree roots may be the least expensive to construct but the most expensive to maintain based on an increase in N concentration with order.

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.

Effects of induction treatments on flowering in Populus deltoides.

Stimulation of early flowering is required to shorten breeding cycles of eastern cottonwood (Populus deltoides Bartr. ex Marsh. var. deltoides), a commercially important and fast-growing hardwood species. A series of experiments was conducted to evaluate the influence of various treatments on flowering in rooted cuttings from mature and juvenile trees. A combined treatment of water stress, root pruning and paclobutrazol was applied to 3-month-old rooted cuttings from mature trees. These cuttings had been subjected to root restriction and long days. All treated plants flowered, whereas no untreated plants formed flower buds. One-year-old rooted cuttings from juvenile trees did not flower when treated with either paclobutrazol, paclobutrazol plus water stress, paclobutrazol plus root pruning, or paclobutrazol plus girdling. This was true both under continuous or periodic growth. Assessment of the lack of flowering in juvenile trees may require an integrated approach that investigates environmental or physiological stimuli, assimilate shift, gibberellic acid type and concentration, and flowering-time gene activity in the new shoots of mature and juvenile cottonwood trees.

Elevated CO2 response of photosynthesis depends on ozone concentration in aspen.

The effect of elevated CO(2) and O(3) on apparent quantum yield (varphi), maximum photosynthesis (P(max)), carboxylation efficiency (V(cmax)) and electron transport capacity (J(max)) at different canopy locations was studied in two aspen (Populus tremuloides) clones of contrasting O(3) tolerance. Local light climate at every leaf was characterized as fraction of above-canopy photosynthetic photon flux density (%PPFD). Elevated CO(2) alone did not affect varphi or P(max), and increased J(max) in the O(3)-sensitive, but not in the O(3)-tolerant clone. Elevated O(3) decreased leaf chlorophyll content and all photosynthetic parameters, particularly in the lower canopy, and the negative impact of O(3) increased through time. Significant interaction effect, whereby the negative impact of elevated O(3) was exaggerated by elevated CO(2) was seen in Chl, N and J(max), and occurred in both O(3)-tolerant and O(3)-sensitive clones. The clonal differences in the level of CO(2)xO(3) interaction suggest a relationship between photosynthetic acclimation and background O(3) concentration.

Will photosynthetic capacity of aspen trees acclimate after long-term exposure to elevated CO2 and O3?

Photosynthetic acclimation under elevated carbon dioxide (CO(2)) and/or ozone (O(3)) has been the topic of discussion in many papers recently. We examined whether or not aspen plants grown under elevated CO(2) and/or O(3) will acclimate after 11 years of exposure at the Aspen Face site in Rhinelander, WI, USA. We studied diurnal patterns of instantaneous photosynthetic measurements as well as A/C(i) measurements monthly during the 2004-2008 growing seasons. Our results suggest that the responses of two aspen clones differing in O(3) sensitivity showed no evidence of photosynthetic and stomatal acclimation under either elevated CO(2), O(3) or CO(2) + O(3). Both clones 42E and 271 did not show photosynthetic nor stomatal acclimation under elevated CO(2) and O(3) after a decade of exposure. We found that the degree of increase or decrease in the photosynthesis and stomatal conductance varied significantly from day to day and from one season to another.

Leaf size and surface characteristics of Betula papyrifera exposed to elevated CO2 and O3.

Betula papyrifera trees were exposed to elevated concentrations of CO(2) (1.4 x ambient), O(3) (1.2 x ambient) or CO(2) + O(3) at the Aspen Free-air CO(2) Enrichment Experiment. The treatment effects on leaf surface characteristics were studied after nine years of tree exposure. CO(2) and O(3) increased epidermal cell size and reduced epidermal cell density but leaf size was not altered. Stomatal density remained unaffected, but stomatal index increased under elevated CO(2). Cuticular ridges and epicuticular wax crystallites were less evident under CO(2) and CO(2) + O(3). The increase in amorphous deposits, particularly under CO(2) + O(3,) was associated with the appearance of elongated plate crystallites in stomatal chambers. Increased proportions of alkyl esters resulted from increased esterification of fatty acids and alcohols under elevated CO(2) + O(3). The combination of elevated CO(2) and O(3) resulted in different responses than expected under exposure to CO(2) or O(3) alone.

Effects of decadal exposure to interacting elevated CO2 and/or O3 on paper birch (Betula papyrifera) reproduction.

We studied the effects of long-term exposure (nine years) of birch (Betula papyrifera) trees to elevated CO(2) and/or O(3) on reproduction and seedling development at the Aspen FACE (Free-Air Carbon Dioxide Enrichment) site in Rhinelander, WI. We found that elevated CO(2) increased both the number of trees that flowered and the quantity of flowers (260% increase in male flower production), increased seed weight, germination rate, and seedling vigor. Elevated O(3) also increased flowering but decreased seed weight and germination rate. In the combination treatment (elevated CO(2)+O(3)) seed weight is decreased (20% reduction) while germination rate was unaffected. The evidence from this study indicates that elevated CO(2) may have a largely positive impact on forest tree reproduction and regeneration while elevated O(3) will likely have a negative impact.

Increases in nitrogen uptake rather than nitrogen-use efficiency support higher rates of temperate forest productivity under elevated CO2.

Forest ecosystems are important sinks for rising concentrations of atmospheric CO(2). In previous research, we showed that net primary production (NPP) increased by 23 +/- 2% when four experimental forests were grown under atmospheric concentrations of CO(2) predicted for the latter half of this century. Because nitrogen (N) availability commonly limits forest productivity, some combination of increased N uptake from the soil and more efficient use of the N already assimilated by trees is necessary to sustain the high rates of forest NPP under free-air CO(2) enrichment (FACE). In this study, experimental evidence demonstrates that the uptake of N increased under elevated CO(2) at the Rhinelander, Duke, and Oak Ridge National Laboratory FACE sites, yet fertilization studies at the Duke and Oak Ridge National Laboratory FACE sites showed that tree growth and forest NPP were strongly limited by N availability. By contrast, nitrogen-use efficiency increased under elevated CO(2) at the POP-EUROFACE site, where fertilization studies showed that N was not limiting to tree growth. Some combination of increasing fine root production, increased rates of soil organic matter decomposition, and increased allocation of carbon (C) to mycorrhizal fungi is likely to account for greater N uptake under elevated CO(2). Regardless of the specific mechanism, this analysis shows that the larger quantities of C entering the below-ground system under elevated CO(2) result in greater N uptake, even in N-limited ecosystems. Biogeochemical models must be reformulated to allow C transfers below ground that result in additional N uptake under elevated CO(2).