Anthropogenic nitrogen deposition ameliorates the decline in tree growth caused by a drier climate.

Most forest ecosystems are simultaneously affected by concurrent global change drivers. However, when assessing these effects, studies have mainly focused on the responses to single factors and have rarely evaluated the joined effects of the multiple aspects of environmental change. Here, we analyzed the combined effects of anthropogenic nitrogen (N) deposition and climatic conditions on the radial growth of Acer saccharum, a dominant tree species in eastern North American forests. We capitalized on a long-term N deposition study, replicated along a latitudinal gradient, that has been taking place for more than 20 yr. We analyzed tree radial growth as a function of anthropogenic N deposition (ambient and experimental addition) and of summer temperature and soil water conditions. Our results reveal that experimental N deposition enhances radial growth of this species, an effect that was accentuated as temperature increased and soil water became more limiting. The spatial and temporal extent of our data also allowed us to assert that the positive effects of growing under the experimental N deposition are likely due to changes in the physiological performance of this species, and not due to the positive correlation between soil N and soil water holding capacity, as has been previously speculated in other studies. Our simulations of tree growth under forecasted climate scenarios specific for this region also revealed that although anthropogenic N deposition may enhance tree growth under a large array of environmental conditions, it will not mitigate the expected effects of growing under the considerably drier conditions characteristic of our most extreme climatic scenario.

Chronic nitrogen deposition influences the chemical dynamics of leaf litter and fine roots during decomposition.

Atmospheric nitrogen deposition induces a forest carbon sink across broad parts of the Northern Hemisphere; this carbon sink may partly result from slower litter decomposition. Although microbial responses to experimental nitrogen deposition have been well-studied, evidence linking these microbial responses to changes in the degradation of specific compounds in decaying litter is sparse. We used wet chemistry and Fourier transform infrared spectroscopy (FTIR) methods to study effects of chronic simulated nitrogen deposition on leaf litter and fine root chemistry during a three-year decomposition experiment at four northern hardwood forests in the north-central USA. Leaf litter and fine roots were highly different in initial chemistry, such as concentrations of acid-insoluble fraction (AIF, or Klason lignin) and condensed tannins (CTs). These initial differences persisted over the course of decomposition. Gravimetrically-defined AIF and lignin/carbohydrate reference IR peak ratios both provide evidence that lignin in fine roots was selectively preserved under simulated nitrogen deposition. Lignin/carbohydrate peak ratios were strongly correlated with AIF, suggesting that AIF is a good predictor of lignin. Because AIF is abundant in fine roots, slower AIF degradation was the major driver of the slower fine root decomposition under nitrogen enrichment, explaining 73.5% of the additional root mass retention. Nitrogen enrichment also slowed the loss of CTs and proteins in fine roots. Nitrogen additions initially slowed the loss of AIF, CTs, and proteins in leaf litter, which was comparatively low in AIF, but these effects disappeared at the later stage and did not affect leaf litter mass loss during the experiment. Our results suggest that decomposition of chemical classes subject to oxidative degradation, such as lignin and CTs, is generally inhibited by nitrogen enrichment; but whether this inhibition eventually slows litter mass loss and leads to organic matter accumulation depends on the initial quantities of these classes in litter.

A dynamic leaf gas-exchange strategy is conserved in woody plants under changing ambient CO2 : evidence from carbon isotope discrimination in paleo and CO2 enrichment studies.

Rising atmospheric [CO2 ], ca , is expected to affect stomatal regulation of leaf gas-exchange of woody plants, thus influencing energy fluxes as well as carbon (C), water, and nutrient cycling of forests. Researchers have proposed various strategies for stomatal regulation of leaf gas-exchange that include maintaining a constant leaf internal [CO2 ], ci , a constant drawdown in CO2 (ca  - ci ), and a constant ci /ca . These strategies can result in drastically different consequences for leaf gas-exchange. The accuracy of Earth systems models depends in part on assumptions about generalizable patterns in leaf gas-exchange responses to varying ca . The concept of optimal stomatal behavior, exemplified by woody plants shifting along a continuum of these strategies, provides a unifying framework for understanding leaf gas-exchange responses to ca . To assess leaf gas-exchange regulation strategies, we analyzed patterns in ci inferred from studies reporting C stable isotope ratios (δ(13) C) or photosynthetic discrimination (∆) in woody angiosperms and gymnosperms that grew across a range of ca spanning at least 100 ppm. Our results suggest that much of the ca -induced changes in ci /ca occurred across ca spanning 200 to 400 ppm. These patterns imply that ca  - ci will eventually approach a constant level at high ca because assimilation rates will reach a maximum and stomatal conductance of each species should be constrained to some minimum level. These analyses are not consistent with canalization toward any single strategy, particularly maintaining a constant ci . Rather, the results are consistent with the existence of a broadly conserved pattern of stomatal optimization in woody angiosperms and gymnosperms. This results in trees being profligate water users at low ca , when additional water loss is small for each unit of C gain, and increasingly water-conservative at high ca , when photosystems are saturated and water loss is large for each unit C gain.

Chronic nitrogen deposition alters tree allometric relationships: implications for biomass production and carbon storage.

As increasing levels of nitrogen (N) deposition impact many terrestrial ecosystems, understanding the potential effects of higher N availability is critical for forecasting tree carbon allocation patterns and thus future forest productivity. Most regional estimates of forest biomass apply allometric equations, with parameters estimated from a limited number of studies, to forest inventory data (i.e., tree diameter). However most of these allometric equations cannot account for potential effects of increased N availability on biomass allocation patterns. Using 18 yr of tree diameter, height, and mortality data collected for a dominant tree species (Acer saccharum) in an atmospheric N deposition experiment, we evaluated how greater N availability affects allometric relationships in this species. After taking into account site and individual variability, our results reveal significant differences in allometric parameters between ambient and experimental N deposition treatments. Large trees under experimental N deposition reached greater heights at a given diameter; moreover, their estimated maximum height (mean ± standard deviation: 33.7 ± 0.38 m) was significantly higher than that estimated under the ambient condition (31.3 ± 0.31 m). Within small tree sizes (5-10 cm diameter) there was greater mortality under experimental N deposition, whereas the relative growth rates of small trees were greater under experimental N deposition. Calculations of stemwood biomass using our parameter estimates for the diameter-height relationship indicated the potential for significant biases in these estimates (~2.5%), with under predictions of stemwood biomass averaging 4 Mg/ha lower if ambient parameters were to be used to estimate stem biomass of trees in the experimental N deposition treatment. As atmospheric N deposition continues to increase into the future, ignoring changes in tree allometry will contribute to the uncertainty associated with aboveground carbon storage estimates across a forest with a large geographic distribution in eastern North America.

Fine roots are the dominant source of recalcitrant plant litter in sugar maple-dominated northern hardwood forests.

Most studies of forest litter dynamics examine the biochemical characteristics and decomposition of leaf litter, but fine roots are also a large source of litter in forests. We quantified the concentrations of eight biochemical fractions and nitrogen (N) in leaf litter and fine roots at four sugar maple (Acer saccharum)-dominated hardwood forests in the north-central United States. We combined these results with litter production data to estimate ecosystem biochemical fluxes to soil. We also compared how leaf litter and fine root biochemistry responded to long-term simulated N deposition. Compared with leaf litter, fine roots contained 2.9-fold higher acid-insoluble fraction (AIF) and 2.3-fold more condensed tannins; both are relatively difficult to decompose. Comparatively, leaf litter had greater quantities of more labile components: nonstructural carbohydrates, cellulose and soluble phenolics. At an ecosystem scale, fine roots contributed over two-thirds of the fluxes of AIF and condensed tannins to soil. Fine root biochemistry was also less responsive than leaf litter to long-term simulated N deposition. Fine roots were the dominant source of difficult-to-decompose plant carbon fractions entering the soil at our four study sites. Based on our synthesis of the literature, this pattern appears to be widespread in boreal and temperate forests.

Redefining fine roots improves understanding of below-ground contributions to terrestrial biosphere processes.

Fine roots acquire essential soil resources and mediate biogeochemical cycling in terrestrial ecosystems. Estimates of carbon and nutrient allocation to build and maintain these structures remain uncertain because of the challenges of consistently measuring and interpreting fine-root systems. Traditionally, fine roots have been defined as all roots ≤ 2 mm in diameter, yet it is now recognized that this approach fails to capture the diversity of form and function observed among fine-root orders. Here, we demonstrate how order-based and functional classification frameworks improve our understanding of dynamic root processes in ecosystems dominated by perennial plants. In these frameworks, fine roots are either separated into individual root orders or functionally defined into a shorter-lived absorptive pool and a longer-lived transport fine-root pool. Using these frameworks, we estimate that fine-root production and turnover represent 22% of terrestrial net primary production globally - a c. 30% reduction from previous estimates assuming a single fine-root pool. Future work developing tools to rapidly differentiate functional fine-root classes, explicit incorporation of mycorrhizal fungi into fine-root studies, and wider adoption of a two-pool approach to model fine roots provide opportunities to better understand below-ground processes in the terrestrial biosphere.

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.

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.

Simulated nitrogen deposition affects community structure of arbuscular mycorrhizal fungi in northern hardwood forests.

Our previous investigation found elevated nitrogen deposition caused declines in abundance of arbuscular mycorrhizal fungi (AMF) associated with forest trees, but little is known about how nitrogen affects the AMF community composition and structure within forest ecosystems. We hypothesized that N deposition would lead to significant changes in the AMF community structure. We studied the diversity and community structure of AMF in northern hardwood forests after more than 12 years of simulated nitrogen deposition. We performed molecular analyses on maple (Acer spp.) roots targeting the 18S rDNA region using the fungal-specific primers AM1 and NS31. PCR products were cloned and identified using restriction fragment length polymorphism (RFLP) and sequencing. N addition significantly altered the AMF community structure, and Glomus group A dominated the AMF community. Some Glomus operational taxonomic units (OTUs) responded negatively to N inputs, whereas other Glomus OTUs and an Acaulospora OTU responded positively to N inputs. The observed effect on community structure implies that AMF species associated with maples differ in their response to elevated nitrogen. Given that functional diversity exists among AMF species and that N deposition has been shown to select less beneficial fungi in some ecosystems, this change in community structure could have implications for the functioning of this type of ecosystem.

Ephemeral root modules in Fraxinus mandshurica.

Historically, ephemeral roots have been equated with 'fine roots' (i.e. all roots of less than an arbitrary diameter, such as 2 mm), but evidence shows that 'fine roots' in woody species are complex branching systems with both rapid-cycling and slow-cycling components. A precise definition of ephemeral roots is therefore needed. Using a branch-order classification, a rhizotron method and sequential sampling of a root cohort, we tested the hypothesis that ephemeral root modules exist within the branching Fraxinus mandshurica (Manchurian ash) root system as distal nonwoody lateral branches, which show anatomical, nutritional and physiological patterns distinct from their woody mother roots. Our results showed that in F. mandshurica, distal nonwoody root branch orders die rapidly as intact lateral branches (or modules). These nonwoody branch orders exhibited highly synchronous changes in tissue nitrogen concentrations and respiration, dominated root turnover, nutrient flux and root respiration, and never underwent secondary development. The ephemeral root modules proposed here may provide a functional basis for differentiating and sampling short-lived absorptive roots in woody plants, and represent a conceptual leap over the traditional coarse-fine root dichotomies based on arbitrary size classes.

Nitrogen turnover in the leaf litter and fine roots of sugar maple.

In order to better understand the nitrogen (N) cycle, a pulse of 15NO3- was applied in 1998 to a sugar maple (Acer saccharum) dominated northern hardwood forest receiving long-term (1994-2008) simulated atmospheric N deposition. Sugar maple leaf litter and live fine-root 15N were quantified for four years prior to labeling and for 11 subsequent years. Continuous sampling of 15N following addition of the tracer enabled calculation of leaf litter and fine-root N pool turnover utilizing an exponential decay function. Fine-root 15N recovery peaked at 3.7% +/- 1.7% the year the tracer was applied, while leaf litter 15N recovery peaked in the two years following tracer application at approximately 8%. These results suggest shoots are primarily constructed from N taken up in previous years, while fine roots are constructed from new N. The residence time of N was 6.5 years in leaf litter and 3.1 years in fine roots. The longer residence time and higher recovery rate are evidence that leaves were a stronger sink for labeled N than fine roots, but the relatively short residence time of tracer N in both pools suggests that there is not tight intra-ecosystem cycling of N in this mature forest.

Reduction of soil carbon formation by tropospheric ozone under increased carbon dioxide levels.

In the Northern Hemisphere, ozone levels in the troposphere have increased by 35 per cent over the past century, with detrimental impacts on forest and agricultural productivity, even when forest productivity has been stimulated by increased carbon dioxide levels. In addition to reducing productivity, increased tropospheric ozone levels could alter terrestrial carbon cycling by lowering the quantity and quality of carbon inputs to soils. However, the influence of elevated ozone levels on soil carbon formation and decomposition are unknown. Here we examine the effects of elevated ozone levels on the formation rates of total and decay-resistant acid-insoluble soil carbon under conditions of elevated carbon dioxide levels in experimental aspen (Populus tremuloides) stands and mixed aspen-birch (Betula papyrifera) stands. With ambient concentrations of ozone and carbon dioxide both raised by 50 per cent, we find that the formation rates of total and acid-insoluble soil carbon are reduced by 50 per cent relative to the amounts entering the soil when the forests were exposed to increased carbon dioxide alone. Our results suggest that, in a world with elevated atmospheric carbon dioxide concentrations, global-scale reductions in plant productivity due to elevated ozone levels will also lower soil carbon formation rates significantly.

Wood {delta}13C, {delta}18O and radial growth responses of residual red pine to variable retention harvesting.

Variable retention harvests are used to enhance the development of structural complexity in managed forests by retaining living trees and other structural legacies from the pre-harvest ecosystem. While harvesting should increase resource availability to residual trees, greater crown exposure may also increase environmental stress, which makes it difficult to predict growth in different structural environments. We used stable carbon isotope ratios (delta(13)C) of annual rings from red pine trees (Pinus resinosa Ait.) as an index of intrinsic water use efficiency (iWUE), the ratio of photosynthetic carbon assimilation (A) to stomatal conductance (g(s)), to better understand how differences in physiological performance relate to growth responses following harvests that left residuals dispersed, aggregated between small (0.1 ha) gaps or aggregated between large (0.3 ha) gaps. Stable oxygen isotope ratios (delta(18)O) were used as an index of g(s) to investigate the drivers behind changes in iWUE. Retention harvesting did not appear to affect delta(13)C or delta(18)O at the stand scale when compared to unharvested control stands, but there was a significant, negative correlation between residual tree delta(13)C and plot basal area in the second and third years after harvesting that suggests declining iWUE as overstory competition increases. Residual tree delta(18)O was similar across treatments and basal areas. Trees in variable retention harvests showed small but positive increases in radial growth from the pre-treatment to post-treatment measurement periods, while radial growth declined in unharvested control stands. There were no significant differences in radial growth among retention treatments. Our results suggest residual red pine in relatively open environments benefit from greater A but do not show evidence of changes in g(s) that would indicate altered water relations.

Species-specific responses to atmospheric carbon dioxide and tropospheric ozone mediate changes in soil carbon.

We repeatedly sampled the surface mineral soil (0-20 cm depth) in three northern temperate forest communities over an 11-year experimental fumigation to understand the effects of elevated carbon dioxide (CO(2)) and/or elevated phyto-toxic ozone (O(3)) on soil carbon (C). After 11 years, there was no significant main effect of CO(2) or O(3) on soil C. However, within the community containing only aspen (Populus tremuloides Michx.), elevated CO(2) caused a significant decrease in soil C content. Together with the observations of increased litter inputs, this result strongly suggests accelerated decomposition under elevated CO(2.) In addition, an initial reduction in the formation of new (fumigation-derived) soil C by O(3) under elevated CO(2) proved to be only a temporary effect, mirroring trends in fine root biomass. Our results contradict predictions of increased soil C under elevated CO(2) and decreased soil C under elevated O(3) and should be considered in models simulating the effects of Earth's altered atmosphere.

Are basidiomycete laccase gene abundance and composition related to reduced lignolytic activity under elevated atmospheric NO3(-) deposition in a northern hardwood forest?

Anthropogenic release of biologically available N has increased atmospheric N deposition in forest ecosystems, which may slow decomposition by reducing the lignolytic activity of white-rot fungi. We investigated the potential for atmospheric N deposition to reduce the abundance and alter the composition of lignolytic basidiomycetes in a regional network of four northern hardwood forest stands receiving experimental NO(3)(-) deposition (30 kg NO(3)(-)-N ha(-1) year(-1)) for a decade. To estimate the abundance of basidiomycetes with lignolytic potential, we used PCR primers targeting laccase (polyphenol oxidase) and quantitative fluorescence PCR to estimate gene copy number. Natural variation in laccase gene size permitted use of length heterogeneity PCR to profile basidiomycete community composition across two sampling dates in forest floor and mineral soil. Although past work has identified significant and consistent negative effects of NO(3)(-) deposition on lignolytic enzyme activity, microbial biomass, soil respiration, and decomposition rate, we found no consistent effect of NO(3)(-) deposition on basidiomycete laccase gene abundance or community profile. Rather, laccase abundance under NO(3)(-) deposition was lower (-52%), higher (+223%), or unchanged, depending on stand. Only a single stand exhibited a significant change in basidiomycete laccase gene profile. Basidiomycete laccase genes occurring in mineral soil were a subset of the genes observed in the forest floor. Moreover, significant effects on laccase abundance were confined to the forest floor, suggesting that species composition plays some role in determining how lignolytic basidiomycetes are affected by N deposition. Community profiles differed between July and October sampling dates, and basidiomycete communities sampled in October had lower laccase gene abundance in the forest floor, but higher laccase abundance in mineral soil. Although experimental N deposition significantly suppresses lignolytic activity in these forests, this change is not related to the abundance or community composition of basidiomycete fungi with laccase genes. Understanding the expression of laccases and other lignolytic enzymes by basidiomycete fungi and other lignin-decaying organisms appears to hold promise for explaining the consistent decline in lignolytic activity elicited by experimental N deposition.

Leaf and canopy conductance in aspen and aspen-birch forests under free-air enrichment of carbon dioxide and ozone.

Increasing concentrations of atmospheric carbon dioxide (CO2) and tropospheric ozone (O3) have the potential to affect tree physiology and structure, and hence forest feedbacks on climate. Here, we investigated how elevated concentrations of CO2 (+45%) and O3 (+35%), alone and in combination, affected conductance for mass transfer at the leaf and canopy levels in pure aspen (Populus tremuloides Michx.) and in mixed aspen and birch (Betula papyrifera Marsh.) forests in the free-air CO2-O3 enrichment experiment near Rhinelander, Wisconsin (Aspen FACE). The study was conducted during two growing seasons, when steady-state leaf area index (L) had been reached after > 6 years of exposure to CO2- and O3-enrichment treatments. Canopy conductance (g(c)) was estimated from stand sap flux, while leaf-level conductance of sun leaves in the upper canopy was derived by three different and independent methods: sap flux and L in combination with vertical canopy modelling, leaf 13C discrimination methodology in combination with photosynthesis modelling and leaf-level gas exchange. Regardless of the method used, the mean values of leaf-level conductance were higher in trees growing under elevated CO2 and/or O3 than in trees growing in control plots, causing a CO2 x O3 interaction that was statistically significant (P < or = 0.10) for sap flux- and (for birch) 13C-derived leaf conductance. Canopy conductance was significantly increased by elevated CO2 but not significantly affected by elevated O3. Investigation of a short-term gap in CO2 enrichment demonstrated a +10% effect of transient exposure of elevated CO2-grown trees to ambient CO2 on g(c). All treatment effects were similar in pure aspen and mixed aspen-birch communities. These results demonstrate that short-term primary stomatal closure responses to elevated CO2 and O3 were completely offset by long-term cumulative effects of these trace gases on tree and stand structure in determining canopy- and leaf-level conductance in pure aspen and mixed aspen-birch forests. Our results, together with the findings from other long-term FACE experiments with trees, suggest that model assumptions of large reductions in stomatal conductance under rising atmospheric CO2 are very uncertain for forests.

Scaling of respiration to nitrogen in leaves, stems and roots of higher land plants.

Using a database of 2510 measurements from 287 species, we assessed whether general relationships exist between mass-based dark respiration rate and nitrogen concentration for stems and roots, and if they do, whether they are similar to those for leaves. The results demonstrate strong respiration-nitrogen scaling relationships for all observations and for data averaged by species; for roots, stems and leaves examined separately; and for life-forms (woody, herbaceous plants) and phylogenetic groups (angiosperms, gymnosperms) considered separately. No consistent differences in the slopes of these log-log scaling relations were observed among organs or among plant groups, but respiration rates at any common nitrogen concentration were consistently lower on average in leaves than in stems or roots, indicating that organ-specific relationships should be used in models that simulate respiration based on tissue nitrogen concentrations. The results demonstrate both common and divergent aspects of tissue-level respiration-nitrogen scaling for leaves, stems and roots across higher land plants, which are important in their own right and for their utility in modelling carbon fluxes at local to global scales.

Soil respiration, root biomass, and root turnover following long-term exposure of northern forests to elevated atmospheric CO2 and tropospheric O3.

The Rhinelander free-air CO(2) enrichment (FACE) experiment is designed to understand ecosystem response to elevated atmospheric carbon dioxide (+CO(2)) and elevated tropospheric ozone (+O(3)). The objectives of this study were: to understand how soil respiration responded to the experimental treatments; to determine whether fine-root biomass was correlated to rates of soil respiration; and to measure rates of fine-root turnover in aspen (Populus tremuloides) forests and determine whether root turnover might be driving patterns in soil respiration. Soil respiration was measured, root biomass was determined, and estimates of root production, mortality and biomass turnover were made. Soil respiration was greatest in the +CO(2) and +CO(2) +O(3) treatments across all three plant communities. Soil respiration was correlated with increases in fine-root biomass. In the aspen community, annual fine-root production and mortality (g m(-2)) were positively affected by +O(3). After 10 yr of exposure, +CO(2) +O(3)-induced increases in belowground carbon allocation suggest that the positive effects of elevated CO(2) on belowground net primary productivity (NPP) may not be offset by negative effects of O(3). For the aspen community, fine-root biomass is actually stimulated by +O(3), and especially +CO(2) +O(3).

delta13C and delta18O trends across overstory environments in whole foliage and cellulose of three Pinus species.

Stable isotope ratios of carbon (delta(13)C) and oxygen (delta(18)O) are increasingly used to investigate environmental influences on plant physiology. Cellulose is often isolated for isotopic studies, but some authors have questioned the value of this process. We studied trends in delta(13)C and delta(18)O of whole foliage and holocellulose from seedlings of three Pinus species across three overstory environments to evaluate the benefits of holocellulose extraction in the context of a traditional ecological experiment. Both tissue types showed increasing delta(13)C from closed-canopy controls to thinned plots to 0.3 ha canopy gaps, and no change in delta(18)O between overstory environments. delta(13)C of P. resinosa and P. strobus was greater than delta(13)C of P. banksiana in whole foliage and holocellulose samples, and there were no differences in delta(18)O associated with species in either tissue type. Our results suggest whole foliage and holocellulose provide similar information about isotopic trends across broad environmental gradients and between species, but holocellulose may be better suited for studying differences in stable isotope composition between multiple species across several treatments.

Simulated atmospheric NO3- deposition increases soil organic matter by slowing decomposition.

Presently, there is uncertainty regarding the degree to which anthropogenic N deposition will foster C storage in the N-limited forests of the Northern Hemisphere, ecosystems which are globally important sinks for anthropogenic CO2. We constructed organic matter and N budgets for replicate northern hardwood stands (n = 4) that have received ambient (0.7-1.2 g N x m(-2) x yr(-1) and experimental NO3- deposition (ambient plus 3 g NO3(-)-N x m(-2) x yr(-1)) for a decade; we also traced the flow of a 15NO3- pulse over a six-year period. Experimental NO3- deposition had no effect on organic matter or N stored in the standing forest overstory, but it did significantly increase the N concentration (+19%) and N content (+24%) of canopy leaves. In contrast, a decade of experimental NO3- deposition significantly increased amounts of organic matter (+12%) and N (+9%) in forest floor and mineral soil, despite no increase in detritus production. A greater forest floor (Oe/a) mass under experimental NO3- deposition resulted from slower decomposition, which is consistent with previously reported declines in lignolytic activity by microbial communities exposed to experimental NO3- deposition. Tracing 15NO3- revealed that N accumulated in soil organic matter by first flowing through soil microorganisms and plants, and that the shedding of 15N-labeled leaf litter enriched soil organic matter over a six-year duration. Our results demonstrate that atmospheric NO3- deposition exerts a direct and negative effect on microbial activity in this forest ecosystem, slowing the decomposition of aboveground litter and leading to the accumulation of forest floor and soil organic matter. To the best of our knowledge, this mechanism is not represented in the majority of simulation models predicting the influence of anthropogenic N deposition on ecosystem C storage in northern forests.