Responses of stomatal density and carbon isotope composition of sugar maple and yellow birch foliage to N, P and CaSiO3 fertilization.

Stomatal density, stomatal length and carbon isotope composition can all provide insights into environmental controls on photosynthesis and transpiration. Stomatal measurements can be time-consuming; it is therefore wise to consider efficient sampling schemes. Knowing the variance partitioning at different measurement levels (i.e., among stands, plots, trees, leaves and within leaves) can aid in making informed decisions around where to focus sampling effort. In this study, we explored the effects of nitrogen (N), phosphorus (P) and calcium silicate (CaSiO3) addition on stomatal density, length and carbon isotope composition (δ13C) of sugar maple (Acer saccharum Marsh.) and yellow birch (Betula alleghaniensis Britton). We observed a positive but small (8%) increase in stomatal density with P addition and an increase in δ13C with N and CaSiO3 addition in sugar maple, but we did not observe effects of nutrient addition on these characteristics in yellow birch. Variability was highest within leaves and among trees for stomatal density and highest among stomata for stomatal length. To reduce variability and increase chances of detecting treatment differences in stomatal density and length, future protocols should consider pretreatment and repeated measurements of trees over time or measure more trees per plot, increase the number of leaf impressions or standardize their locations, measure more stomata per image and ensure consistent light availability.

Patterns of physical, chemical, and metabolic characteristics of sugar maple leaves with depth in the crown and in response to nitrogen and phosphorus addition.

Few previous studies have described the patterns of leaf characteristics in response to nutrient availability and depth in the crown. Sugar maple has been studied for both sensitivity to light, as a shade-tolerant species, and sensitivity to soil nutrient availability, as a species in decline due to acid rain. To explore leaf characteristics from the top to bottom of the canopy, we collected leaves along a vertical gradient within mature sugar maple crowns in a full-factorial nitrogen (N) by phosphorus (P) addition experiment in three forest stands in central New Hampshire, USA. Thirty-two of the 44 leaf characteristics had significant relationships with depth in the crown, with the effect of depth in the crown strongest for leaf area, photosynthetic pigments and polyamines. Nitrogen addition had a strong impact on the concentration of foliar N, chlorophyll, carotenoids, alanine and glutamate. For several other elements and amino acids, N addition changed patterns with depth in the crown. Phosphorus addition increased foliar P and boron (B); it also caused a steeper increase of P and B with depth in the crown. Since most of these leaf characteristics play a direct or indirect role in photosynthesis, metabolic regulation or cell division, studies that ignore the vertical gradient may not accurately represent whole-canopy performance.

Tree variability limits the detection of nutrient treatment effects on sap flux density in a northern hardwood forest.

The influence of nutrient availability on transpiration is not well understood, in spite of the importance of transpiration to forest water budgets. Soil nutrients have the potential to affect tree water use through indirect effects on leaf area or stomatal conductance. For example, following addition of calcium silicate to a watershed at Hubbard Brook, in New Hampshire, streamflow was reduced for 3 years, which was attributed to a 25% increase in evapotranspiration associated with increased foliar production. The first objective of this study was to quantify the effect of nutrient availability on sap flux density in a nitrogen, phosphorus, and calcium addition experiment in New Hampshire in which tree diameter growth, foliar chemistry, and soil nutrient availability had responded to treatments. We measured sap flux density in American beech (Fagus grandifolia, Ehr.), red maple (Acer rubrum L.), sugar maple (Acer saccharum Marsh.), white birch (Betula papyrifera Marsh.), or yellow birch (Betula alleghaniensis Britton.) trees, over five years of experiments in five stands distributed across three sites. In 2018, 3 years after a calcium silicate addition, sap flux density averaged 36% higher in trees in the treatment than the control plot, but this effect was not very significant (p = 0.07). Our second objective was to determine whether this failure to detect effects with greater statistical confidence was due to small effect sizes or high variability among trees. We found that tree-to-tree variability was high, with coefficients of variation averaging 39% within treatment plots. Depending on the species and year of the study, the minimum difference in sap flux density detectable with our observed variability ranged from 46% to 352%, for a simple ANOVA. We analyzed other studies reported in the literature that compared tree water use among species or treatments and found detectable differences ranging from 16% to 78%. Future sap flux density studies could benefit from power analyses to guide sampling intensity. Including pretreatment data, in the case of manipulative studies, would also increase statistical power.

N and P constrain C in ecosystems under climate change: Role of nutrient redistribution, accumulation, and stoichiometry.

We use the Multiple Element Limitation (MEL) model to examine responses of 12 ecosystems to elevated carbon dioxide (CO2 ), warming, and 20% decreases or increases in precipitation. Ecosystems respond synergistically to elevated CO2 , warming, and decreased precipitation combined because higher water-use efficiency with elevated CO2 and higher fertility with warming compensate for responses to drought. Response to elevated CO2 , warming, and increased precipitation combined is additive. We analyze changes in ecosystem carbon (C) based on four nitrogen (N) and four phosphorus (P) attribution factors: (1) changes in total ecosystem N and P, (2) changes in N and P distribution between vegetation and soil, (3) changes in vegetation C:N and C:P ratios, and (4) changes in soil C:N and C:P ratios. In the combined CO2 and climate change simulations, all ecosystems gain C. The contributions of these four attribution factors to changes in ecosystem C storage varies among ecosystems because of differences in the initial distributions of N and P between vegetation and soil and the openness of the ecosystem N and P cycles. The net transfer of N and P from soil to vegetation dominates the C response of forests. For tundra and grasslands, the C gain is also associated with increased soil C:N and C:P. In ecosystems with symbiotic N fixation, C gains resulted from N accumulation. Because of differences in N versus P cycle openness and the distribution of organic matter between vegetation and soil, changes in the N and P attribution factors do not always parallel one another. Differences among ecosystems in C-nutrient interactions and the amount of woody biomass interact to shape ecosystem C sequestration under simulated global change. We suggest that future studies quantify the openness of the N and P cycles and changes in the distribution of C, N, and P among ecosystem components, which currently limit understanding of nutrient effects on C sequestration and responses to elevated CO2 and climate change.

Foliar nutrient concentrations of six northern hardwood species responded to nitrogen and phosphorus fertilization but did not predict tree growth.

Foliar chemistry can be useful for diagnosing soil nutrient availability and plant nutrient limitation. In northern hardwood forests, foliar responses to nitrogen (N) addition have been more often studied than phosphorus (P) addition, and the interactive effects of N and P addition have rarely been described. In the White Mountains of central New Hampshire, plots in ten forest stands of three age classes across three sites were treated annually beginning in 2011 with 30 kg N ha-1 y-1 or 10 kg P ha-1 y-1 or both or neither-a full factorial design. Green leaves of American beech (Fagus grandifolia Ehrh.), pin cherry (Prunus pensylvanica L.f.), red maple (Acer rubrum L.), sugar maple (A. saccharum Marsh.), white birch (Betula papyrifera Marsh.), and yellow birch (B. alleghaniensis Britton) were sampled pre-treatment and 4-6 years post-treatment in two young stands (last cut between 1988-1990), four mid-aged stands (last cut between 1971-1985) and four mature stands (last cut between 1883-1910). In a factorial analysis of species, stand age class, and nutrient addition, foliar N was 12% higher with N addition (p < 0.001) and foliar P was 45% higher with P addition (p < 0.001). Notably, P addition reduced foliar N concentration by 3% (p = 0.05), and N addition reduced foliar P concentration by 7% (p = 0.002). When both nutrients were added together, foliar P was lower than predicted by the main effects of N and P additions (p = 0.08 for N × P interaction), presumably because addition of N allowed greater use of P for growth. Foliar nutrients did not differ consistently with stand age class (p ≥ 0.11), but tree species differed (p ≤ 0.01), with the pioneer species pin cherry having the highest foliar nutrient concentrations and the greatest responses to nutrient addition. Foliar calcium (Ca) and magnesium (Mg) concentrations, on average, were 10% (p < 0.001) and 5% lower (p = 0.01), respectively, with N addition, but were not affected by P addition (p = 0.35 for Ca and p = 0.93 for Mg). Additions of N and P did not affect foliar potassium (K) concentrations (p = 0.58 for N addition and p = 0.88 for P addition). Pre-treatment foliar N:P ratios were high enough to suggest P limitation, but trees receiving N (p = 0.01), not P (p = 0.64), had higher radial growth rates from 2011 to 2015. The growth response of trees to N or P addition was not explained by pre-treatment foliar N, P, N:P, Ca, Mg, or K.

Length and colonization rates of roots associated with arbuscular or ectomycorrhizal fungi decline differentially with depth in two northern hardwood forests.

Ectomycorrhizal (EM) and arbuscular mycorrhizal (AM) fungi are often studied independently, and thus little is known regarding differences in vertical distribution of these two groups in forests where they co-occur. We sampled roots at two soil depths in two northern hardwood stands in Bartlett, New Hampshire, co-dominated by tree species that associate with AM or EM fungi. Root length of both groups declined with depth. More importantly, root length of EM plant species exceeded that of AM plants at 0-10-cm depth, while AM exceeded EM root length at 30-50-cm depth. Colonization rates were similar between mineral and organic portions of the shallow (0-10 cm) samples for EM and AM fungi and declined dramatically with depth (30-50 cm). The ratio of EM to AM fungal colonization declined with depth, but not as much as the decline in root length with depth, resulting in greater dominance by EM fungi near the surface and by AM fungi at depth. The depth distribution of EM and AM roots may have implications for soil carbon accumulation as well as for the success of the associated tree species.

Quantifying uncertainty in annual runoff due to missing data.

Long-term streamflow datasets inevitably include gaps, which must be filled to allow estimates of runoff and ultimately catchment water budgets. Uncertainty introduced by filling gaps in discharge records is rarely, if ever, reported. We characterized the uncertainty due to streamflow gaps in a reference watershed at the Hubbard Brook Experimental Forest (HBEF) from 1996 to 2009 by simulating artificial gaps of varying duration and flow rate, with the objective of quantifying their contribution to uncertainty in annual streamflow. Gaps were filled using an ensemble of regressions relating discharge from nearby streams, and the predicted flow was compared to the actual flow. Differences between the predicted and actual runoff increased with both gap length and flow rate, averaging 2.8% of the runoff during the gap. At the HBEF, the sum of gaps averaged 22 days per year, with the lowest and highest annual uncertainties due to gaps ranging from 1.5 mm (95% confidence interval surrounding mean runoff) to 21.1 mm. As a percentage of annual runoff, uncertainty due to gap filling ranged from 0.2-2.1%, depending on the year. Uncertainty in annual runoff due to gaps was small at the HBEF, where infilling models are based on multiple similar catchments in close proximity to the catchment of interest. The method demonstrated here can be used to quantify uncertainty due to gaps in any long-term streamflow data set, regardless of the gap-filling model applied.

Determining optimal sampling strategies for monitoring mercury and reproductive success in common loons in the Adirondacks of New York.

The common loon (Gavia immer), a top predator in the freshwater food web, has been recognized as an important bioindicator of aquatic mercury (Hg) pollution. Because capturing loons can be difficult, statistical approaches are needed to evaluate the efficiency of Hg monitoring. Using data from 1998 to 2016 collected in New York's Adirondack Park, we calculated the power to detect temporal changes in loon Hg concentrations and fledging success as a function of sampling intensity. There is a tradeoff between the number of lakes per year and the number of years needed to detect a particular rate of change. For example, a 5% year-1 change in Hg concentration could be detected with a sampling effort of either 15 lakes per year for 10 years, or 5 lakes per year for 15 years, given two loons sampled per lake per year. A 2% year-1 change in fledging success could be detected with a sampling effort of either 40 lakes per year for 15 years, or 30 lakes per year for 20 years. We found that more acidic lakes required greater sampling intensity than less acidic lakes for monitoring Hg concentrations but not for fledging success. Power analysis provides a means to optimize the sampling designs for monitoring loon Hg concentrations and reproductive success. This approach is applicable to other monitoring schemes where cost is an issue.

Spatial patterns and temporal trends in mercury concentrations in common loons (Gavia immer) from 1998 to 2016 in New York's Adirondack Park: has this top predator benefitted from mercury emission controls?

Mercury (Hg), a neurotoxic pollutant, can be transported long distances through the atmosphere and deposited in remote areas, threatening aquatic wildlife through methylation and bioaccumulation. Over the last two decades, air quality management has resulted in decreases in Hg emissions from waste incinerators and coal-fired power plants across North America. The common loon (Gavia immer) is an apex predator of the aquatic food web. Long-term monitoring of Hg in loons can help track biological recovery in response to the declines in atmospheric Hg that have been documented in the northeastern USA. To assess spatial patterns and temporal trends in Hg exposure of the common loon in the Adirondack Park of New York State, we analyzed Hg concentrations in loon blood and egg samples from 116 lakes between 1998 and 2016. We found spatially variable Hg concentrations in adult loon blood and feathers across the Park. Loon Hg concentrations (converted to female loon units) increased 5.7% yr-1 from 1998 to 2010 (p = 0.04), and then stabilized at 1.70 mg kg-1 from 2010 to 2016 (p = 0.91), based on 760 observations. Concentrations of Hg in juvenile loons also increased in the early part of the record, stabilizing 2 years before Hg concentrations stabilized in adults. For 52 individual lakes with samples from at least 4 different years, loon Hg increased in 34 lakes and decreased in 18 lakes. Overall, we found a delayed recovery of Hg concentrations in loons, despite recent declines in atmospheric Hg.

Estimating uncertainty in the volume and carbon storage of downed coarse woody debris.

Downed coarse woody debris, also known as coarse woody detritus or downed dead wood, is challenging to estimate for many reasons, including irregular shapes, multiple stages of decay, and the difficulty of identifying species. In addition, some properties are commonly not measured, such as wood density and carbon concentration. As a result, there have been few previous evaluations of uncertainty in estimates of downed coarse woody debris, which are necessary for analysis and interpretation of the data. To address this shortcoming, we quantified uncertainties in estimates of downed coarse woody debris volume and carbon storage using data collected from permanent forest inventory plots in the northeastern United States by the Forest Inventory and Analysis program of the USDA Forest Service. Quality assurance data collected from blind remeasurement audits were used to quantify error in diameter measurements, hollowness of logs, species identification, and decay class determination. Uncertainty estimates for density, collapse ratio, and carbon concentration were taken from the literature. Estimates of individual sources of uncertainty were combined using Monte Carlo methods. Volume estimates were more reliable than carbon storage, with an average 95% confidence interval of 15.9 m3 /ha across the 79 plots evaluated, which was less than the mean of 31.2 m3 /ha. Estimates of carbon storage (and mass) were more uncertain, due to poorly constrained estimates of the density of wood. For carbon storage, the average 95% confidence interval was 11.1 Mg C/ha, which was larger than the mean of 4.6 Mg C/ha. Accounting for the collapse of dead wood as it decomposes would improve estimates of both volume and carbon storage. On the other hand, our analyses suggest that consideration of the hollowness of downed coarse woody debris pieces could be eliminated in this region, with little effect. This study demonstrates how uncertainty analysis can be used to quantify confidence in estimates and to help identify where best to allocate resources to improve monitoring designs.

Downsizing a long-term precipitation network: Using a quantitative approach to inform difficult decisions.

The design of a precipitation monitoring network must balance the demand for accurate estimates with the resources needed to build and maintain the network. If there are changes in the objectives of the monitoring or the availability of resources, network designs should be adjusted. At the Hubbard Brook Experimental Forest in New Hampshire, USA, precipitation has been monitored with a network established in 1955 that has grown to 23 gauges distributed across nine small catchments. This high sampling intensity allowed us to simulate reduced sampling schemes and thereby evaluate the effect of decommissioning gauges on the quality of precipitation estimates. We considered all possible scenarios of sampling intensity for the catchments on the south-facing slope (2047 combinations) and the north-facing slope (4095 combinations), from the current scenario with 11 or 12 gauges to only 1 gauge remaining. Gauge scenarios differed by as much as 6.0% from the best estimate (based on all the gauges), depending on the catchment, but 95% of the scenarios gave estimates within 2% of the long-term average annual precipitation. The insensitivity of precipitation estimates and the catchment fluxes that depend on them under many reduced monitoring scenarios allowed us to base our reduction decision on other factors such as technician safety, the time required for monitoring, and co-location with other hydrometeorological measurements (snow, air temperature). At Hubbard Brook, precipitation gauges could be reduced from 23 to 10 with a change of <2% in the long-term precipitation estimates. The decision-making approach illustrated in this case study is applicable to the redesign of monitoring networks when reduction of effort seems warranted.

Concentrations and content of mercury in bark, wood, and leaves in hardwoods and conifers in four forested sites in the northeastern USA.

Mercury (Hg) is deposited from the atmosphere to remote areas such as forests, but the amount of Hg in trees is not well known. To determine the importance of Hg in trees, we analyzed foliage, bark and bole wood of eight tree species at four sites in the northeastern USA (Huntington Forest, NY; Sleepers River, VT; Hubbard Brook, NH; Bear Brook, ME). Foliar concentrations of Hg averaged 16.3 ng g-1 among the hardwood species, which was significantly lower than values in conifers, which averaged 28.6 ng g-1 (p < 0.001). Similarly, bark concentrations of Hg were lower (p < 0.001) in hardwoods (7.7 ng g-1) than conifers (22.5 ng g-1). For wood, concentrations of Hg were higher in yellow birch (2.1-2.8 ng g-1) and white pine (2.3 ng g-1) than in the other species, which averaged 1.4 ng g-1 (p < 0.0001). Sites differed significantly in Hg concentrations of foliage and bark (p = 0.02), which are directly exposed to the atmosphere, but the concentration of Hg in wood depended more on species (p < 0.001) than site (p = 0.60). The Hg contents of tree tissues in hardwood stands, estimated from modeled biomass and measured concentrations at each site, were higher in bark (mean of 0.10 g ha-1) and wood (0.16 g ha-1) than in foliage (0.06 g ha-1). In conifer stands, because foliar concentrations were higher, the foliar pool tended to be more important. Quantifying Hg in tree tissues is essential to understanding the pools and fluxes of Hg in forest ecosystems.

Phosphorus limitation of aboveground production in northern hardwood forests.

Forest productivity on glacially derived soils with weatherable phosphorus (P) is expected to be limited by nitrogen (N), according to theories of long-term ecosystem development. However, recent studies and model simulations based on resource optimization theory indicate that productivity can be co-limited by N and P. We conducted a full factorial N × P fertilization experiment in 13 northern hardwood forest stands of three age classes in central New Hampshire, USA, to test the hypothesis that forest productivity is co-limited by N and P. We also asked whether the response of productivity to N and P addition differs among species and whether differential species responses contribute to community-level co-limitation. Plots in each stand were fertilized with 30 kg N·ha-1 ·yr-1 , 10 kg P·ha-1 ·yr-1 , N + P, or neither nutrient (control) for four growing seasons. The productivity response to treatments was assessed using per-tree annual relative basal area increment (RBAI) as an index of growth. RBAI responded significantly to P (P = 0.02) but not to N (P = 0.73). However, evidence for P limitation was not uniform among stands. RBAI responded to P fertilization in mid-age (P = 0.02) and mature (P = 0.07) stands, each taken as a group, but was greatest in N-fertilized plots of two stands in these age classes, and there was no significant effect of P in the young stands. Both white birch (Betula papyrifera Marsh.) and beech (Fagus grandifolia Ehrh.) responded significantly to P; no species responded significantly to N. We did not find evidence for N and P co-limitation of tree growth. The response to N + P did not differ from that to P alone, and there was no significant N × P interaction (P = 0.68). Our P limitation results support neither the N limitation prediction of ecosystem theory nor the N and P co-limitation prediction of resource optimization theory, but could be a consequence of long-term anthropogenic N deposition in these forests. Inconsistencies in response to P suggest that successional status and variation in site conditions influence patterns of nutrient limitation and recycling across the northern hardwood forest landscape.

Soil nitrogen affects phosphorus recycling: foliar resorption and plant-soil feedbacks in a northern hardwood forest.

Previous studies have attempted to link foliar resorption of nitrogen and phosphorus to their. respective availabilities in soil, with mixed results. Based on resource optimization theory, we hypothesized that the foliar resorption of one element could be driven by the availability of another element. We tested various measures of soil N and P as predictors of N and P resorption in six tree species in 18 plots across six stands at the Bartlett Experimental Forest, New Hampshire, USA. Phosphorus resorption efficiency (P < 0.01) and proficiency (P = 0.01) increased with soil N content. to 30 cm depth, suggesting that trees conserve P based on the availability of soil N. Phosphorus resorption also increased with soil P content, which is difficult to explain basdd on single-element limitation, butfollows from the correlation between soil N and soil P. The expected single-element relationships were evident only in the 0 horizon: P resorption was high where resin-available P was low in the Oe (P < 0.01 for efficiency, P < 0.001 for proficiency) and N resorption was high where potential N mineralization in the Oa was low (P < 0.01 for efficiency and 0.11 for proficiency). Since leaf litter is a principal source of N and P to the 0 horizon, low nutrient availability there could be a result rather than a cause of high resorption. The striking effect of soil N content on foliar P resorption is the first evidence of multiple-element control on nutrient resorption to be reported from an unmanipulated ecosystem.

From missing source to missing sink: long-term changes in the nitrogen budget of a northern hardwood forest.

Biogeochemical monitoring for 45 years at the Hubbard Brook Experimental Forest in New Hampshire has revealed multiple surprises, seeming contradictions, and unresolved questions in the long-term record of ecosystem nitrogen dynamics. From 1965 to 1977, more N was accumulating in living biomass than was deposited from the atmosphere; the "missing" N source was attributed to biological fixation. Since 1992, biomass accumulation has been negligible or even negative, and streamwater export of dissolved inorganic N has decreased from ~4 to ~1 kg of N ha(-1) year(-1), despite chronically elevated atmospheric N deposition (~7 kg of N ha(-1) year(-1)) and predictions of N saturation. Here we show that the ecosystem has shifted to a net N sink, either storing or denitrifying ~8 kg of N ha(-1) year(-1). Repeated sampling over 25 years shows that the forest floor is not detectably accumulating N, but the C:N ratio is increasing. Mineral soil N has decreased nonsignificantly in recent decades, but the variability of these measurements prevents detection of a change of <700 kg of N ha(-1). Whether the excess N is accumulating in the ecosystem or lost through denitrification will be difficult to determine, but the distinction has important implications for the local ecosystem and global climate.

Lead reduction and redistribution in the forest floor in New Hampshire northern hardwoods.

Because of the affinity of organic matter for lead, atmospheric loadings of this pollutant have been strongly retained in the forest floor. With the regulation of Pb emissions, loadings have decreased. We measured changes in Pb in forest floor horizons at a variety of northern hardwood sites in New Hampshire from the late 1970s to the 1990s. In all seven of the sites in which horizons were distinguished within the forest floor, Pb was found to be declining in the upper (Oie) horizon, but not in the underlying Oa and A horizons. At the Hubbard Brook Experimental Forest (HBEF), this loss from the Oie resulted in a 36% loss of Pb from the forest floor as a whole between 1976 and 1997 (p < 0.001). In contrast, in six stands in the Bartlett Experimental Forest (BEF), losses of Pb averaging >50% from the Oi and Oe horizons (p = 0.01) between 1979 and 1994 were compensated by gains in the Oa and A horizons. Similarly, at seven additional stands in the White Mountain National Forest, changes in the forest floor as a whole from 1980 to 1995 were not statistically significant (redistribution within the forest floor was not evaluated at these sites). Lead concentrations were highest in the Oe or Oie in the 1970s, but were highest in the Oa horizon in the 1990s. There was no significant pattern of Pb loss or retention as a function of stand age across all the sites.

Modeling changes in red spruce carbon balance and allocation in response to interacting ozone and nutrient stresses.

The simulation model TREGRO was developed to analyze the response of red spruce saplings to multiple stresses, such as drought, nutrient deficiency, and exposure to pollutants. The model provides a method of identifying changes in structural and non-structural carbon resources in the tree that may become measurable only after many years of exposure. The model is based on the assumption that the ability of plants to take up and use carbon, water, and nutrients depends on the interrelationships in availability among the three resources. Consequently, the model simulates the simultaneous cycling of these resources. In the model, the tree is divided into the following compartments: a canopy of leaves grouped by age class, branches, stem, and coarse and fine roots in a number of soil horizons. In each of these compartments we track three carbon pools: living structure, dead structure or wood, and total non-structural carbohydrate. The model calculates the photosynthesis of an entire red spruce tree each hour as a function of ambient environmental conditions and the availability of light, water, and nutrients; the daily redistribution of carbon throughout the plant; and the loss of carbon by respiration and senescence. To accomplish this task, the model tracks the flow of carbon dioxide to the sites of fixation within the leaves, the availability of light in the canopy, water and nutrient resources in each of three soil horizons, and the amounts of these resources taken up by the tree. Soil and plant water potentials, photosynthesis, and leaf respiration are simulated on an hourly timestep; nutrient uptake, allocation and growth are computed on a daily timestep. Through a set of example simulations, we demonstrate how the model can be used to examine the mechanisms by which plants respond to stresses experienced alone and in combination. The model was used to predict the growth decrease and the shifting pattern of carbon allocation expected for an isolated tree exposed to ozone and decreased nutrient availability due to acidic deposition. Decreased nutrient availability resulted in decreased growth and preferential carbon allocation to roots, which helped to alleviate the nutrient stress. Ozone stress also resulted in decreased plant growth but had the opposite effect on allocation patterns, with most of the growth reduction occurring in roots. The effect of simultaneous ozone and nutrient stress on tree growth was less than the sum of the independent single stresses, contrary to our expectation. This modeling approach can aid in evaluating the long-term effect of stress on resource availability, the potential for gradual deterioration of tree health under long periods of stress, and imbalances in growth accompanying shifts in carbon allocation caused by stress.