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The ability of woody plants to survive for decades, centuries, sometimes millennia, is due in part to their capacity to withstand environmental stress by shifting their resources from roots, to shoots, to storage reserves.

Two important processes which may limit productivity gains in forest ecosystems with rising atmospheric CO

SUMMARY Fine root demography was quantified in response to patches of increased water and nitrogen availability in a natural, second-growth, mixed hardwood forest in northern Michigan, USA. As expected, the addition of water and water plus nitrogen resulted in a significant overall increase in the production of new fine roots. New root production was much greater in response to water plus nitrogen when compared with water alone, and the duration of new root production was related to the length of resource addition in the water plus nitrogen treatments; the average difference in new root length between the 20 vs. 40 d additions of water plus nitrogen amounted to almost 600%. Roots produced in response to the additions of water and water plus nitrogen lived longer than roots in the control treatments. Thus, additions of water and water plus nitrogen influenced both the proliferation of new roots and their longevity, with both proliferation and longevity related to the type and duration of resource supply. Results suggest that root longevity and mortality may be plastic in response to changes in soil resource availability, as is well known for root proliferation.

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) CO

Atmospheric nitrogen deposition increases forest carbon sequestration across broad parts of the Northern Hemisphere. Slower organic matter decomposition and greater soil carbon accumulation could contribute to this increase in carbon sequestration. We investigated the effects of chronic simulated nitrogen deposition on leaf litter and fine root decomposition at four sugar maple (Acer saccharum)- dominated northern hardwood forests. At these sites, we previously observed that nitrogen additions increased soil organic carbon and altered litter chemistry. We conducted a 3-year decomposition study with litter bags. Litter production of leaves and fine roots were combined with decomposition dynamics to estimate how fine roots and leaf litter contribute to soil organic carbon. We found that nitrogen additions marginally stimulated early-stage decomposition of leaf litter, an effect associated with previously documented changes in litter chemistry. In contrast, nitrogen additions inhibited the later stages of fine root decomposition, which is consistent with observed decreases in lignin-degrading enzyme activities with nitrogen additions at these sites. At the ecosystem scale, slower fine root decomposition led to additional root mass retention (g m-2), and this greater retention of root residues was estimated to explain 5-51% of previously documented carbon accumulation in the surface soil due to nitrogen additions. Our results demonstrated that simulated nitrogen deposition created contrasting effects on the decomposition of leaf litter and fine roots. Although previous nitrogen deposition studies have focused on leaf litter, this work suggests that slower fine root decomposition is a major driver of soil organic carbon accumulation under elevated nitrogen deposition.

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.

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.

Summary � 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.

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.

The storage and flow of carbon (C) into and out of forests can differ under the influence of dominant tree species because of species-based variation in C production, decomposition, retention, and harvest-based export. Following abandonment of agricultural activities in the first half of the 20th century, many landscapes of the Great Lakes region (USA) were planted to red pine (Pinus resinosa) or naturally regenerated to northern hardwood species including sugar maple (Acer saccharum), red oak (Quercus rubra) and red maple (Acer rubrum). We located eight pairs of adjacent, similarly aged (∼60 yr) stands of planted red pine and naturally regenerated hardwood forests on previous agricultural fields. We found that the hardwood forests stored more C than pine stands (255 vs. 201 Mg C ha−1), with both storing substantially more than an adjacent area maintained as pasture (107 Mg C ha−1). The greater accumulation of C in the hardwood stands occurred mostly in living biomass. No significant differences for soil C (to 1 m depth) were found between forest types, despite significantly higher belowground inputs and aboveground litterfall in hardwood stands. Notably, both forest types had about 18% more soil C than the pasture, with O horizon C accounting for about one-third of the increase under trees. Forest type had no significant effect on estimated amount of exported C despite fairly large differences in projected end uses (solid wood products, land-fills, bioenergy). Using adjacent pasture as our baseline condition, we combined estimated on-site accumulation rates with estimates of exported C, and found that average total C sequestration rates were higher for hardwood (2.9 Mg C ha−1 yr−1) than red pine plots (2.3 Mg C ha−1 yr−1). The modeled potential contribution of exported C to these sequestration rate estimates did not differ between species, but the fate of modeled post-harvest off-site C may exert a large influence on sequestration rate estimates depending on actual displacement actions, including product longevity. These results show that tree species selection has the potential to impact C sequestration rates but effects vary by ecosystem component and could not be predicted from previous species effects studies.

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.

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

We tested the hypothesis that elevated CO

Humans have altered the global nitrogen (N) cycle, greatly increasing atmospheric nitrogen deposition in industrialized regions of the world. Groundcover plants can be sensitive indicators of nitrogen deposition impacts. Here, we report results from repeated measurements over a 7 year period of groundcover (plants 50% (P

Industrialization has greatly affected the biogeochemistry of northern forests by increasing the atmospheric deposition of acid and nitrogen (N). In 1990, the US Congress amended the Clean Air Act to include tighter emissions regulations; this reduced acid deposition (by >50% in this study), but did not effectively lower N deposition. Here, we demonstrate that since this legislation was enacted, there have been marked decreases in sulfur (−16%), calcium (−17%), and aluminum (−42%) concentrations in sugar maple (Acer saccharum) foliage across the Upper Great Lakes region of the US, signaling a declining influence of acid deposition. In contrast, N deposition has persistently been over 75% greater than the amount of N needed to offset annual plant N sequestration, creating increases in N availability and soil N leaching. Recent emissions regulations will reduce N deposition somewhat, but further increases in soil N availability and leaching are likely. Policy decisions regarding N deposition will have to weigh increased carbon storage against negative impacts on water quality and species diversity.

In most forested landscapes, the organization of plant communities across stream valleys is thought to be regulated by a complex set of interactions including flooding, landform properties, and vegetation. However, few studies have directly examined the relative influence of frequent and infrequent flooding, as well as landform properties, on riparian plant community organization in moderately or deeply entrenched stream valleys where the magnitude and extent of frequent flooding may be constrained by local stream valley characteristics. Our approach, which we applied in an old-growth northern hardwood watershed, integrated detailed plant community surveys with a GIS and watershed surface hydrology model that allowed us to model water surface elevation associated with different flood magnitudes and recurrence intervals for specific locations across the old-growth watershed. Our results show that irrespective of stream valley geomorphology, the ground-flora exhibits a high rate of species replacement across the stream valley at low elevations, which are the most susceptible to frequent and more extreme infrequent flooding. However, over 50 % of the major shifts in ground-flora community composition and almost all of the shifts in overstory composition occur beyond the direct influence of flooding, especially in the high-gradient moderately and deeply entrenched stream valleys. In these areas, landform boundaries and changes in the environmental properties associated with these boundaries appear to be the primary factors controlling changes in vegetation across the stream valley.

Recent meta-analyses of experimental studies simulating increased anthropogenic nitrogen (N) deposition in forests reveal greater soil carbon (C) storage under elevated levels of atmospheric N deposition. However, these effects have not yet been included in ecosystem-scale models of soil C and N cycling and it is unclear whether increased soil C storage results from slower decomposition rates or a reduced extent of decomposition (for example, an increase in the amount of litter entering slowly decaying humus pools). To test these alternatives, we conducted a meta-analysis of litter decomposition data. We then used the results from our meta-analysis to model C and N cycling in four sugar maple forests in Michigan using an ecosystem process model (TRACE). We compared model results testing our alternative hypotheses to field data on soil C storage from a 17-year N deposition experiment. Using data from published litter decomposition studies in forests, we determined that, on average, exogenous N inputs decreased lignin decomposition rates by 30% and increased cellulose decomposition by 9%. In the same set of litter decomposition studies increased exogenous N availability increased the amount of litter entering slowly decaying humus pools in a manner significantly related to the lignocellulose index of decaying litter. Incorporating changes to decomposition rates in TRACE did not accurately reproduce greater soil C storage observed in our field study with experimentally elevated N deposition. However, when changes in the extent of decomposition were incorporated in TRACE, the model produced increased soil C storage by increasing the amount of litter entering the humus pool and accurately represented C storage in plant and soil pools under experimental N deposition. Our modeling results and meta-analysis indicate that the extent of litter decay as humus is formed, rather than slower rates of litter decay, is likely responsible for the accumulation of organic matter, and hence soil C storage, under experimental N deposition. This effect should be incorporated in regional to global-scale models simulating the C balance of forest ecosystems in regions receiving elevated N deposition.

Plant growth responses to rising atmospheric CO2 and O3 vary among genotypes and between species, which could plausibly influence the strength of competitive interactions for soil N. Ascribable to the size-symmetric nature of belowground competition, we reasoned that differential growth responses to CO2 and O3 should shift as juvenile individuals mature, thereby altering competitive hierarchies and forest composition. In a 12-year-long forest FACE experiment, we used tracer 15 N and whole-plant N content to assess belowground competitive interactions among five Populus tremuloides genotypes, between a single P. tremuloides genotype and Betula papryrifera, as well as between the same single P. tremuloides genotype and Acer saccharum. Under elevated CO2, the amount of soil N and 15 N obtained by the P. tremuloides genotype common to each community was contingent on the nature of belowground competition. When this genotype competed with its congeners, it obtained equivalent amounts of soil N and tracer 15 N under ambient and elevated CO2; however, its acquisition of soil N under elevated CO2 increased by a significant margin when grown in competition with B. papyrifera (+30%) and A. saccharum (+60%). In contrast, elevated O3 had no effect on soil N and 15 N acquisition by the P. tremuloides genotype common in each community, regardless of competitive interactions. Under elevated CO2, the rank order of N acquisition among P. tremuloides genotypes shifted over time, indicating that growth responses to CO2 change during ontogeny; this was not the case under elevated O3 .I n the aspen-birch community, the competitive advantage elevated CO2 initially conveyed on birch diminished over time, whereas maple was a poor competitor for soil N in all regards. The extent to which elevated CO2 and O3 will shape the genetic structure and composition of future forests is, in part, contingent on the time-dependent effects of belowground competition on plant growth response.

Summary 1. During the next century, atmospheric nitrogen (N) deposition is projected to more than double, potentially leading to a decline in plant diversity as well as a change in plant community composition and structure. 2. In a decade-long field experiment, simulated atmospheric N deposition has slowed litter decay, resulting in an accumulation of forest floor (i.e. Oi & Oe horizons). We reasoned that a greater forest floor mass under simulated N deposition would impose a physical barrier to sugar maple Acer saccharum seedling establishment, thereby reducing seedling populations of an ecologically and economically important tree species. 3. To test this idea, we first quantified sugar maple seedling abundance in replicate northern hardwood forest stands receiving ambient atmospheric N (7–12 kg N ha )1 year )1 ) and experimental atmospheric N deposition, simulating future amounts in eastern North America (ambient plus 30 kg NO3 Nh a )1 year )1 ). Then, we experimentally manipulated forest floor mass under ambient and simulated N deposition treatments. Finally, we transplanted first-year established seedlings into areas receiving ambient and simulated N deposition and quantified their mortality after 1 year. 4. First-year seedling abundance did not differ under ambient and simulated N deposition; however, there were greater abundances of second- and third-to-fifth-year seedlings under ambient N deposition ( P< 0AE001). In all cases, experimental manipulation to increase forest floor mass, equivalent to that under simulated N deposition, resulted in significantly (P =0 AE001) fewer established individuals, regardless of whether the greater forest floor mass occurred under ambient or simulated N deposition. Finally, fewer 1-year-old transplanted seedlings survived when grown under simulated N, albeit that result was not statistically significant. 5. Synthesis and applications. The slowing of decay and the accumulation of forest floor under anthropogenic N deposition can negatively impact seedling survival and potentially alter stand development and structural diversity. As atmospheric N deposition increases globally, it becomes necessary to understand the mechanisms that lead to population changes for ecologically important tree species. The responses we document should be considered in simulations of future of forest dynamics, as atmospheric N deposition continues to increase, specifically when sugar maple life-history traits are included to simulate regeneration, structural diversity and stand development.

In this review, we present a conceptual model which links plant communities and saprotrophic microbial communities through the reciprocal exchange of growth-limiting resources. We discuss the numerous ways human-induced environmental change has directly and indirectly impacted this relationship, and review microbial responses that have occurred to date. We argue that compositional shifts in saprotrophic microbial communities underlie functional responses to environmental change that have ecosystem-level implications. Drawing on a long-term, large-scale, field experiment, we illustrate how and why chronic atmospheric N deposition can alter saprotrophic communities in the soil of a wide-spread sugar maple (Acer saccharum) ecosystem in northeastern North America, resulting in the slowing of plant litter decay, the rapid accumulation of soil organic matter, and the accelerated production and loss of dissolved organic carbon (DOC). Compositional shifts in soil microbial communities, mediated by ecological interactions among soil saprotrophs, appear to lie at the biogeochemical heart of ecosystem response to environmental change.

Elevated concentrations of atmospheric CO2 and tropospheric O3 will profoundly influence future forest productivity, but our understanding of these influences over the long-term is poor. Leaves are key indicators of productivity and we measured the mass, area, and nitrogen concentration of leaves collected in litter traps from 2002 to 2008 in three young northern temperate forest communities exposed to elevated CO2 and/or elevated O3 since 1998. On average, the overall effect of elevated CO2 (+CO2 and +CO2+O3 versus ambient and +O3) was to increase leaf mass by 36% whereas the overall effect of elevated O3 was to decrease leaf mass by 13%, with similar effects on stand leaf area. However, there were important CO2 × O3 × year interactions wherein some treatment effects on leaf mass changed dramatically relative to ambient from 2002 to 2008. For example, stimulation by the +CO2 treatment decreased (from +52 to +25%), whereas the deleterious effects of the +O3 treatment increased (from −5 to −18%). In comparison, leaf mass in the +CO2+O3 treatment was similar to ambient throughout the study. Forest composition influenced these responses: effects of the +O3 treatment on community-level leaf mass ranged from +2 to −19%. These findings are evidence that community composition, stand development processes, CO2, and O3 strongly interact. Changes in leaf nitrogen concentration were inconsistent, but leaf nitrogen mass (g m−2) was increased by elevated CO2 (+30%) and reduced by elevated O3 (−16%), consistent with observations that nitrogen cycling is accelerated by elevated CO2 but retarded by elevated O3.

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.

Specific root respiration rates typically increase with increasing tissue N concentration. As a result, it is often assumed that external factors inducing greater root N concentration, such as chronic N deposition, will lead to increased respiration rates. However, enhanced N availability also alters root biomass, making the ecosystem-level consequences on whole-root-system respiration uncertain. The objective of this study was to determine the effects of chronic experimental N deposition on root N concentrations, specific respiration rates, and biomass for four northern hardwood forests in Michigan. Three of the six measurement plots at each location have received experimental N deposition (3 g NO 3 -N m 2 yr 1 ) since 1994. We measured specific root respiration rates and N concentrations of roots from four size classes (

Several genes in the aspen genome have been modified to generate stem wood with lower lignin content and an altered lignin composition. Lower lignin in wood reduces the time and energy required for pulping. Further, this modification can also increase the allocation of photosynthate to cellulose and total biomass production, potentially increasing CO 2 -sequestration capacity. However, widespread planting of trees with altered lignin content and composition could alter soil organic-C dynamics in complex ways. To further examine the effects of altered lignin biosynthesis on plant growth and accrual of soil organic C (SOC), we conducted a repeated greenhouse study with four lines of transgenic aspen (Populus tremuloides Michx.) and one wild-type (control) aspen. Accrual of aspen-derived SOC was quantified by growing aspen trees (C3 plants) in C4 soil and measuring changes in the natural abundance of d13C. We measured plant growth, biomass, and C content and combined these data with SOC measurements to create C budgets for the plant mesocosms. Lignin modifications resulted in differences in the accrual of aspen-derived SOC and total mesocosm C, primarily due to differences in biomass between genetically modified lines of aspen. One genetic alteration (low lignin, line 23) was able to perform similarly or better than the wild-type aspen (control, line 271) without altering SOC. Alterations in lignin structure (S : G ratios) had negative effects on biomass production and SOC formation. The addition of new (aspen-derived) SOC was proportional to the loss of existing SOC, evidence for a priming effect. The pool of new SOC was related to total plant biomass, suggesting that the effects of lignin modification on SOC are driven by changes in plant growth.

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.

Responses of mature trees to chronic N additions are poorly understood in ecosystems with high seasonal and spatial variability. To determine the effects of increased N deposition on mature conifers, we fertilized a pinon-juniper woodland in New Mexico at a rate equivalent to the urban interface. Fertilization (10 g m −2 y −1 ) reduced numbers of mycorrhizae and increased leaf production in the ectomycorrhizal (EM) pinon but not in arbuscular mycorrhizal (AM) juniper. Based on N fractionation between EM fungal sporocarps and pinon, EM in pinon utilized 20% of the net primary production in control plots. No sporocarps were produced in fertilized plots. N uptake by pinon could be accounted for by fertilization without mycorrhizae. Leaf N and size increased with fertilization in both species, and positively correlated with leaf δ 13 C. Leaf N:P increased in pinon but not juniper. Pinon mortality commenced in the N-fertilized plots in 2001, a year before the widespread die-off in western conifers, and continued through 2003. No mortality was observed in control plots or in junipers. The coupling of N enrichment and mycorrhizal decline could affect pinon production and mortality in semi-arid woodlands in the western US.

Increased nitrogen (N) deposition caused by human activities has altered ecosystem functioning and biodiversity. To understand the effects of altered N availability, we measured the abundance of arbuscular mycorrhizal fungi (AMF) and the microbial community in northern hardwood forests exposed to long-term (12 years) simulated N deposition (30 kg N ha -1 y -1 ) using phospholipid

Plant and microbial use of nitrogen (N) can be simultaneously mutualistic and competitive, particularly in ecosystems dominated by mycorrhizal fungi. Our goal was to quantify plant uptake of organic and inorganic N across a broad latitudinal gradient of forest ecosystems that varied with respect to overstory taxon, edaphic characteristics, and dominant mycorrhizal association. Using 13C and 15N, we observed in situ the cycling dynamics of NH4 + and glycine through various soil pools and fine roots over 14 days. Recovery of 15N as soil N varied with respect to N form, forest type, and sampling period; however, there were similarities in the cycling dynamics of glycine and NH4 + among all forest types. Microbial immobilization of 15N was immediately apparent for both treatments and represented the largest sink (~25%) for 15N among extractable soil N pools during the first 24 h. In contrast, fine roots were a relatively small sink (

Free amino acids (FAA) constitute a significant fraction of dissolved organic nitrogen (N) in forest soils and play an important role in the N cycle of these ecosystems. However, comparatively little attention has been given to their role as labile carbon (C) substrates that might influence the metabolic status of resident microbial populations. We hypothesized that the residence time of simple C substrates, such as FAA, are mechanistically linked to the turnover of endogenous soil C pools. We tested this hypothesis across a latitudinal gradient of forested ecosystems that differ sharply with regard to climate, overstory taxon, and edaphic properties. Using a combined laboratory and field approach, we compared the turnover of isotopically labeled glycine in situ to the turnover of mineralizable soil C (Cmin) at each site. The turnover of glycine was rapid (residence times

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.

Overstory conditions influence understory microclimate and resource availability, leading to gradients in evaporative demand and moisture availability that influence seedling water relations. Partial canopies may either reduce seedling moisture stress by ameliorating environmental conditions, or increase moisture stress by reducing soil moisture availability. This study used stable isotope ratios of oxygen (δ 18 O) and carbon (δ 13 C) and mass-based foliar nitrogen concentrations to investigate changes in transpiration ( E ), stomatal conductance ( g s ) and intrinsic water use efficiency (iWUE) of pine seedlings across an overstory gradient from open canopy gap environments to closed canopy forest. Foliar δ 18 O increased sharply from basal areas of 0–10 m 2 ha −1 in Pinus banksiana , Pinus resinosa , and Pinus strobus seedlings, followed by a more gradual increase with further increases in basal area. Foliar δ 13 C followed a similar, but less pronounced pattern in P. banksiana and P. strobus seedlings, and had no apparent relationship with overstory basal area in P. resinosa seedlings. The slope of the δ 18 O:δ 13 C relationship was positive for every species. Foliar nitrogen concentrations were not correlated with overstory basal area. These results suggest seedling E declined as overstory basal area increased due to reductions in g s , while iWUE increased slightly from open gaps to partial canopy environments. Open gap environments appear to provide sufficient moisture to sustain high leaf-level gas exchange rates in the species we studied, while relatively small increases in overstory basal area apparently promote rapid declines in g s , leading to greatly reduced seedling water loss and small increases in iWUE.

Variable retention harvest systems are encouraged to promote complexity in managed forests, and aggregated retention has been suggested as a means of reducing moisture stress in residual trees. We studied the impacts of within-aggregate position on growth and foliar physiology to better understand the spatial dynamics of residual-tree responses to aggregated retention harvests in even-aged Pinus resinosa Ait. stands. Distance from edge and edge aspect influenced radial growth, volume increment, and growth efficiency, but only edge aspect affected foliar nitrogen content. Spatial variables had no significant relationships with foliar carbon isotope ratios (δ13C). Increases in radial growth, volume increment, and growth efficiency following harvesting were greatest near edges and in the northeastern quadrants of aggregates that received mechanical understory release treatments, and lowest in the southeastern quadrant of aggregates and near aggregate centers. Foliar nitrogen content was highest in the southwestern quadrants of aggregates that received understory release treatments, and lowest in the northwestern quadrants of aggregates. Our results suggest spatial relationships are important determinants of residual-tree responses to aggregated retention harvests.

Gradients of light and moisture availability peak at different positions within canopy gaps in northern latitudes providing the opportunity for niche partitioning in and around gaps based on differences in individual species’ life history attributes. This gap partitioning offers potential for increasing diversity in forests impacted by gap-creating disturbances. We examined resource availability and the physiological performance of three Pinus species with varying tolerances for shade and moisture stress across large (0.3 ha) canopy gaps to investigate relationships between gap position and species performance. Light availability was lowest in southern gap edges, while water availability was lowest in northern edges, and higher at gap interior positions than edges. Pinus banksiana seedlings had higher light-saturated CO2 assimilation rates than P. resinosa or P. strobus seedlings at interior gap positions, and outperformed P. strobus at northern gap edges, but there were no differences between species at southern edges. Both transpiration and stomatal conductance were greatest for P. banksiana in gap centers, but showed few differences between species at edges. Foliar nitrogen concentrations were highest for P. banksiana, suggesting the dominance of this species in central gap locations may be due to a combination of high photosynthetic capacity and tight stomatal control to regulate moisture stress at drier gap positions. Our results suggest P. banksiana seedlings may be competitively superior in gap positions with high light and moisture availability, but P. resinosa and P. strobus become competitive under the drier conditions and moderate shade near gap edges. These findings support the concept of gap partitioning, and suggest silvicultural systems that incorporate patch cuttings could be used to promote diverse regeneration in northern pine forests.

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.

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.

Plants influence soil carbon (C) formation through the quality and quantity of C released to soil. Soil type, in turn can modify a plant's influence on soil through effects on plant production, tissue quality and regulation of soil C decomposition and stabilization. Wild-type aspen and three transgenic aspen lines expressing reduced stem lignin concentrations and/or increased syringyl (S) to guaiacyl (G) ratio lignin were grown in greenhouse mesocosms containing a sandy loam, a silt loam, or a clay loam soil for 6 months in order to examine the effects of altered lignin biosynthesis and soil type on biomass partitioning (above vs. belowground) and soil C processes. Results indicated that soil type significantly affected plant performance. Aspen grown in soils with high sand/ low clay content accumulated the most total biomass, while aspen grown in soils with high clay content accumulated the least total biomass. These reductions in growth combined with specific soil characteristics led to differences among soil types in soil C formation. Transformed aspen expressing high syringyl/guaiacyl (S/G) lignin accumulated less total plant C and subsequently accumulated less aspen derived C in soil. Reduced lignin content alone in aspen did not affect plant growth or soil C formation. There were significant soil type × genetic line interactions indicating that growth and soil C formation for transgenic and wild type aspen lines varied among the different soil types. Given these interactions, future investigation needs to include long-term field studies across a range of soil types before transgenic aspen are widely planted.

The ability of an ecosystem to retain anthropogenic nitrogen (N) deposition is dependent upon plant and soil sinks for N, the strengths of which may be altered by chronic atmospheric N deposition. Sugar maple (Acer saccharum Marsh.), the dominant overstory tree in northern hardwood forests of the Lake States region, has a limited capacity to take up and assimilate NO 3 − . However, it is uncertain whether long-term exposure to NO 3 − deposition might induce NO 3 − uptake by this ecologically important overstory tree. Here, we investigate whether 10 years of experimental NO 3 − deposition (30 kg N ha−1 y−1) could induce NO 3 − uptake and assimilation in overstory sugar maple (approximately 90 years old), which would enable this species to function as a direct sink for atmospheric NO 3 − deposition. Kinetic parameters for NH 4 + and NO 3 − uptake in fine roots, as well as leaf and root NO 3 − reductase activity, were measured under conditions of ambient and experimental NO 3 − deposition in four sugar maple-dominated stands spanning the geographic distribution of northern hardwood forests in the Upper Lake States. Chronic NO 3 − deposition did not alter the V max or K m for NO 3 − and NH 4 + uptake nor did it influence NO 3 − reductase activity in leaves and fine roots. Moreover, the mean V max for NH 4 + uptake (5.15 μmol 15N g−1 h−1) was eight times greater than the V max for NO 3 − uptake (0.63 μmol 15N g−1 h−1), indicating a much greater physiological capacity for NH 4 + uptake in this species. Additionally, NO 3 − reductase activity was lower than most values for woody plants previously reported in the literature, further indicating a low physiological potential for NO 3 − assimilation in sugar maple. Our results demonstrate that chronic NO 3 − deposition has not induced the physiological capacity for NO 3 − uptake and assimilation by sugar maple, making this dominant species an unlikely direct sink for anthropogenic NO 3 − deposition.

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 towards 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. This article is protected by copyright. All rights reserved. Rising atmospheric [CO2], c(a), 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], c(i), a constant drawdown in CO2 (c(a)-c(i)), and a constant c(i)/c(a). 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 c(a). 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 c(a). To assess leaf gas-exchange regulation strategies, we analyzed patterns in c(i) inferred from studies reporting C stable isotope ratios (C-13) or photosynthetic discrimination () in woody angiosperms and gymnosperms that grew across a range of c(a) spanning at least 100ppm. Our results suggest that much of the c(a)-induced changes in c(i)/c(a) occurred across c(a) spanning 200 to 400ppm. These patterns imply that c(a)-c(i) will eventually approach a constant level at high c(a) 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 c(i). 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 c(a), when additional water loss is small for each unit of C gain, and increasingly water-conservative at high c(a), when photosystems are saturated and water loss is large for each unit C gain.

Concentrations of soluble soil sugars, soluble phenolic acids, and free amino acids were measured in three forest communities at the FACTS-II Aspen FACE Site near Rhinelander, WI, in order to better understand how elevated atmospheric CO2 and O3 are influencing soil nutrient availability and cycling. Sugars, phenolic acids, and amino acids are mostly derived from plant and microbial processes, and have the potential to be influenced by changes in carbon inputs. We hypothesized that concentrations in the soil would parallel increases seen in biological activity, due to greater net primary productivity under elevated CO2 and seasonal patterns of root growth. Chemical analysis of soils revealed marginally significant increases of total soluble sugars and total soluble phenolic acids in the elevated CO2 treatment (+27 mg kg � 1 , +0.02mmol g � 1 ), but there were no significant differences in concentrations due to elevated O3 or CO2+O3. Total free amino acid concentrations were not affected by any of the treatments, but significant shifts in individual amino acids were observed. Elevated CO2 and the interaction treatment (elevated CO2+O3) increased aspartic acid concentrations, while elevated O3 treatment decreased the concentration of valine. Concentrations of sugars increased throughout the growing season, while phenolic acids were constant and amino acids decreased. The birch–aspen community had the highest concentration of phenolic acids and sugars overall, while maple–aspen had the lowest. These findings suggest that concentrations of soluble sugars, soluble phenolic acids, and free amino acids in the soil are strongly influenced by soil properties, plant and microbial activity, plant community composition, and to a lesser degree, changes in atmospheric CO2 and O3. r 2007 Elsevier Ltd. All rights reserved.

Anthropogenic O3 and CO2-induced declines in soil N availability could counteract greater plant growth in a CO2-enriched atmosphere, thereby reducing net primary productivity (NPP) and the potential of terrestrial ecosystems to sequester anthropogenic CO2. Presently, it is uncertain how increasing atmospheric CO2 and O3 will alter plant N demand and the acquisition of soil N by plants as well as the microbial supply of N from soil organic matter. To address this uncertainty, we initiated an ecosystem-level 15N tracer experiment at the Rhinelander (Wisconsin, USA) free air CO2-O3 enrichment (FACE) facility to understand how projected increases in atmospheric CO2 and 03 alter the distribution and flow of N in developing northern temperate forests. Tracer amounts of 15NH4+ were applied to the forest floor of developing Populus tremuloides and P. tremuloides-Betula papyrifera communities that have been exposed to factorial CO2 and O3 treatments for seven years. One year after isotope addition, both forest communities exposed to elevated CO2 obtained greater amounts of 15N (29%) and N (40%) from soil, despite no change in soil N availability or plant N-use efficiency. As such, elevated CO2 increased the ability of plants to exploit soil for N, through the development of a larger root system. Conversely, elevated O3 decreased the amount of 15N (-15%) and N (-29%) in both communities, a response resulting from lower rates of photosynthesis, decreases in growth, and smaller root systems that acquired less soil N. Neither CO2 nor 03 altered the amount of N or 15N recovery in the forest floor, microbial biomass, or soil organic matter. Moreover, we observed no interaction between CO2 and 03 on the amount of N or 15N in any ecosystem pool, suggesting that 03 could exert a negative effect regardless of CO2 concentration. In a CO2-enriched atmosphere, greater belowground growth and a more thorough exploitation of soil for growth-limiting N is an important mechanism sustaining the enhancement of NPP in developing forests (0-8 years following establishment). However, as CO2 accumulates in the Earth's atmosphere, future O3 concentrations threaten to diminish the enhancement of plant growth, decrease plant N acquisition, and lessen the storage of anthropogenic C in temperate forests.

Research aimed at understanding how the global carbon balance will change with elevated CO(2) has largely ignored the responses of individual species and genotypes. Yet, plant traits strongly influence the biogeochemical cycling of carbon. Here, we illustrate how differences in inter- and intraspecific responses to elevated CO(2) affect not only physiology and growth, but also higher order biotic interactions and lifetime fitness, ultimately leading to new ecosystem assemblages. We assert that the unique combination of inter- and intraspecific traits in these ecosystem assemblages ultimately determine how ecosystems respond to elevated atmospheric CO(2). Thus, the identity of species and genotypes in an ecosystem is a crucial element to consider in forecasts of global carbon balance.

As human activity continues to increase CO2 and O3, broad expanses of north temperate forests will be simultaneously exposed to elevated concentrations of these trace gases. Although both CO2 and O3 are potent modifiers of plant growth, we do not understand the extent to which they alter competition for limiting soil nutrients, like nitrogen (N). We quantified the acquisition of soil N in two 8-year-old communities composed of trembling aspen genotypes (n 55) and trembling aspen‐paper birch which were exposed to factorial combinations of CO2 (ambient and 560lLL � 1 ) and O3 (ambient 530‐40 vs. 50‐60nLL � 1 ). Tracer amount of 15 NH4 1 were applied to soil to determine how these trace gases altered the competitive ability of genotypes and species to acquire soil N. One year after isotope addition, we assessed N acquisition by measuring the amount of 15 N tracer contained in the plant canopy (i.e. recent N acquisition), as well as the total amount of canopy N (i.e. cumulative N acquisition). Exposure to elevated CO2 differentially altered recent and cumulative N acquisition among aspen genotypes, changing the rank order in which they obtained soil N. Elevated O3 also altered the rank order in which aspen genotypes obtained soil N by eliciting increases, decreases and no response among genotypes. If aspen genotypes respond similarly under field conditions, then rising concentrations of CO2 and O3 could alter the structure of aspen populations. In the aspen‐birch community, elevated CO2 increased recent N (i.e. 15 N) acquisition in birch (68%) to a greater extent than aspen (19%), suggesting that, over the course of this experiment, birch had gained a competitive advantage over aspen. The response of genotypes and species to rising CO2 and O3 concentrations, and how these responses are modified by competitive interactions, has the potential to change the future composition and productivity of northern temperate forests.

A simple system based on web-FACE technology was designed and implemented as an approach to label plant-assimilated carbon (C) with 13C. The labeling system avoids the use of a chamber or other enclosure, instead distributing CO2 heavily enriched in 13C at near atmospheric concentrations to the tree foliage through the use of porous tubing. The system was applied to three plantation grown juvenile larch (Larix spp.) trees during the daylight hours over the course of five days in the middle of the growing season. Relative to control trees, fumigation with enriched CO2 resulted in significantly 13C-enriched foliar respiration and nighttime soil respiration. Enrichment was also created in the foliar tissue, but differences between labeled and control trees were not statistically significant. Temporal and spatial variation in the strength of the isotopic label did occur, and modifications to the system are suggested to limit the variation. The approach should enable the implementation of pulse-chase experiments designed to understand plant source-sink relationships or experiments designed to understand the flux of C from plant roots into the soil food web.

Sugar maple (Acer saccharum Marsh.)-dominated northern hardwood forests of the Great Lakes Region commonly receive elevated levels of atmospheric nitrate (NO 3 − ) deposition, which can alter belowground carbon (C) cycling. Past research has demonstrated that chronic experimental NO 3 − deposition (3 g N m−2 y−1 above ambient) elicits a threefold increase in the leaching loss of dissolved organic carbon (DOC). Here, we used DOC collected from tension-cup lysimeters to test whether increased DOC export under experimental NO 3 − deposition originated from forest floor or mineral soil organic matter (SOM). We used DOC radiocarbon dating to quantify C sources and colorimetric assays to measure DOC aromaticity and soluble polyphenolic content. Our results demonstrated that DOC exports are primarily derived from new C (

The development and use of transgenic plants has steadily increased, but there are still little data about the responses of soil microorganisms to these genetic modifications. We utilized a greenhouse trial approach to evaluate the effects of altered stem lignin in trembling aspen (Populus tremuloides) on soil microbial communities in three soils which differed in their chemical and physical properties; they included a sandy loam (CO-Colorado), a silt loam (KS-Kansas), and a clay loam (TX-Texas). Three transgenic aspen lines were developed from a natural clone common to the Great Lakes region of North America. The concentrations of stem lignin concentrations were reduced by 35% (Line 23), 40% (Line 141) and 50% (Line 72). Line 72 and Line 141 also had a 40 and 20% increase in syringyl-type stem lignin, respectively. Indirectly, these modifications resulted in increased (5–13%) and decreased (−5 to −57%) levels of root production across the lines and soil types. Responses of the soil microbial communities were investigated using: phospholipid fatty acids (PLFA), neutral lipid fatty acids (NLFA), and 3) extracellular enzyme assays. PLFA analyses indicated that there were large differences in microbial community composition between the three soils. Similarly, there were large differences in total NLFA between soils, with the KS soils having the highest amount and CO the lowest. Enzyme activities did not differ between soils, except for cellubiohydrolase, which was highest in CO soil. Across all three soils, responses to the four genetic lines were not consistent. Interactions between soil type and genetic line make it difficult to assess the potential ecological impacts of transgenic aspen on soil microbial communities and their associated functions. Given these interactions, field trials with transgenic aspen should encompass the wide range of soils targeted for commercial planting in order to determine their effect(s) on the resident soil microbial community.

Carbon (C) allocation to the perennial coarse-root system of trees contributes to ecosystem C sequestration through formation of long-lived live wood biomass and, following senescence, by providing a large source of nutrient- poor detrital C. Our understanding of the controls on C allocation to coarse-root growth is rudimentary, but it has im- portant implications .for projecting belowground net primary production responses to global change. Age-related changes in C allocation to coarse roots represent a critical uncertainty for modeling landscape-scale C storage and cy- cling. We used a 55 year chronosequence approach with complete above- and below-ground harvests to assess the ef- fects of stand development on biomass partitioning in red pine (Pinus resinosa Ait.), a commercially important pine species. Averaged within sita, individual-tree rootlshoot ratios were dynamic across stand development, changing from 0.17 at 2-, 3-, and 5-year-old sites, to 0.80 at the 8-year-old site, to 0.29 at the 55-year-old site. The results of our study suggest that a current research challenge is to determine the generality of patterns of root-shoot biomass parti- ' tioning through stand development for both coniferous and hardwood forest types, and to document how these patterns change as a function of stand age, tree size, environment, and management. RCsumC : L'allocation du carbone (C) aux grosses racines ptrennes des arbres contribue au pikgeage du C des Ccosys- tkmes en produisant une biomasse vivante de bois d'une grande longiviti et en creant, aprbs la senescence, une impor- tante source de C dans des dktritus i faible teneur en nutriments. Cependant, notre comprthension des facteurs qui contrelent l'allocation du C B la croissance des grosses racines est rudimentaire, ce qui enbdne d'importantes rkpercus- sions pour la prediction de la production primaire nette des racines en reaction aux changements climatiques. Les chan- gements dans l'allocation du C aux grosses racines en fonction de l'gge reprCsentent une incertitude cruciale pour modtliser I'entreposage et le recyclage du C B l'kchelle du paysage. Nous avons eu recours 2 une chronosBquence d'une durke de 55 ans, qui incluait la rtcolte compMte des tissus a6riens et racinaires, pour ivaluer les effets du dive

We conducted a glasshouse mesocosm study that combined (13)C isotope techniques with wild-type and transgenic aspen (Populus tremuloides) in order to examine how altered lignin biosynthesis affects plant production and soil carbon formation. Our transgenic aspen lines expressed low stem lignin concentration but normal cellulose concentration, low lignin stem concentration with high cellulose concentration or an increased stem syringyl to guaiacyl lignin ratio. Large differences in stem lignin concentration observed across lines were not observed in leaves or fine roots. Nonetheless, low lignin lines accumulated 15-17% less root C and 33-43% less new soil C than the control line. Compared with the control line, transformed aspen expressing high syringyl lignin accumulated 30% less total plant C - a result of greatly reduced total leaf area - and 70% less new soil C. These findings suggest that altered stem lignin biosynthesis in Populus may have little effect on the chemistry of fine roots or leaves, but can still have large effects on plant growth, biomass partitioning and soil C formation.