Alison H. Magill

Nitrogen Saturation in Temperate Forest Ecosystems

N itrogen emissions to the atmosphere due to human activity remain elevated in industrialized regions of the world and are accelerating in many developing regions (Galloway 1995). Although the deposition of sulfur has been reduced over much of the United States and Europe by aggressive environmental protection policies, current nitrogen deposition reduction targets in the US are modest. Nitrogen deposition remains relatively constant in the northeastern United States and is increasing in the Southeast and the West (Fenn et al. in press). The US acid deposition effects

Is Nitrogen Deposition Altering the Nitrogen Status of Northeastern Forests

Abstract Concern is resurfacing in the United States over the long-term effects of excess nitrogen (N) deposition and mobility in the environment. We present here a new synthesis of existing data sets for the northeastern United States, intended to answer a single question: Is N deposition altering the N status of forest ecosystems in this region? Surface water data suggest a significant increase in nitrate losses with N deposition. Soil data show an increase in nitrification with decreasing ratio of soil carbon to nitrogen (C:N) but weaker relationships between N deposition and soil C:N ratio or nitrification. Relationships between foliar chemistry and N deposition are no stronger than with gradients of climate and elevation. The differences in patterns for these three groups of indicators are explained by the degree of spatial and temporal integration represented by each sample type. The surface water data integrate more effectively over space than the foliar or soil data and therefore allow a more comprehensive view of N saturation. We conclude from these data that N deposition is altering N status in northeastern forests.

Vertical transport of dissolved organic C and N under long-term N amendments in pine and hardwood forests

At the Harvard Forest, Massachusetts, a long-term effort is under way to study responses in ecosystem biogeochemistry to chronic inputs of N in atmospheric deposition in the region. Since 1988, experimental additions of NH4NO3 (0, 5 and 15 g N m−2 yr−1) have been made in two forest stands:Pinus resinosa (red pine) and mixed hardwood. In the seventh year of the study, we measured solute concentrations and estimated solute fluxes in throughfall and at two soil depths, beneath the forest floors (Oa) and beneath the B horizons.Beneath the Oa, concentrations and fluxes of dissolved organic C and N (DOC and DON) were higher in the coniferous stand than in the hardwood stand. The mineral soil exerted a strong homogenizing effect on concentrations beneath the B horizons. In reference plots (no N additions), DON composed 56% (pine) and 67% (hardwood) of the total dissolved nitrogen (TDN) transported downward from the forest floor to the mineral soil, and 98% of the TDN exported from the solums. Under N amendments, fluxes of DON from the forest floor correlated positively with rates of N addition, but fluxes of inorganic N from the Oa exceeded those of DON. Export of DON from the solums appeared unaffected by 7 years of N amendments, but as in the Oa, DON composed smaller fractions of TDN exports under N amendments. DOC fluxes were not strongly related to N amendment rates, but ratios of DOC:DON often decreased.The hardwood forest floor exhibited a much stronger sink for inorganic N than did the pine forest floor, making the inputs of dissolved N to mineral soil much greater in the pine stand. Under the high-N treatment, exports of inorganic N from the solum of the pine stand were increased >500-fold over reference (5.2 vs. 0.01 g N m−2 yr−1), consistent with other manifestations of nitrogen saturation. Exports of N from the solum in the pine forest decreased in the order NO3-N> NH4-N> DON, with exports of inorganic N 14-fold higher than exports of DON. In the hardwood forest, in contrast, increased sinks for inorganic N under N amendments resulted in exports of inorganic N that remained lower than DON exports in N-amended plots as well as the reference plot.

Long-Term Nitrogen Additions and Nitrogen Saturation in Two Temperate Forests

ABSTRACT This article reports responses of two different forest ecosystems to 9 years (1988–96) of chronic nitrogen (N) additions at the Harvard Forest, Petersham, Massachusetts. Ammonium nitrate (NH4NO3) was applied to a pine plantation and a native deciduous broad-leaved (hardwood) forest in six equal monthly doses (May–September) at four rates: control (no fertilizer addition), low N (5 g N m-2 y-1), high N (15 g N m-2y-1), and low N + sulfur (5 g N m-2 y-1 plus 7.4 g S m-2 y-1). Measurements were made of net N mineralization, net nitrification, N retention, wood production, foliar N content and litter production, soil C and N content, and concentrations of dissolved organic carbon (DOC) and nitrogen (DON) in soil water. In the pine stand, nitrate losses were measured after the first year of additions (1989) in the high N plot and increased again in 1995 and 1996. The hardwood stand showed no significant increases in nitrate leaching until 1995 (high N only), with further increases in 1996. Overall N retention efficiency (percentage of added N retained) over the 9-year period was 97–100% in the control and low N plots of both stands, 96% in the hardwood high N plot, and 85% in the pine high N plot. Storage in aboveground biomass, fine roots, and soil extractable pools accounted for only 16–32% of the added N retained in the amended plots, suggesting that the one major unmeasured pool, soil organic matter, contains the remaining 68–84%. Short-term redistribution of 15N tracer at natural abundance levels showed similar division between plant and soil pools. Direct measurements of changes in total soil C and N pools were inconclusive due to high variation in both stands. Woody biomass production increased in the hardwood high N plot but was significantly reduced in the pine high N plot, relative to controls. A drought-induced increase in foliar litterfall in the pine stand in 1995 is one possible factor leading to a measured increase in N mineralization, nitrification, and nitrate loss in the pine high N plot in 1996.

Plant and Soil Responses to Chronic Nitrogen Additions at the Harvard Forest, Massachusetts

Data are presented on changes in plant and soil processes in two forest types (red pine plantation and oak-maple forest) at the Harvard Forest, Petersham, Massachusetts, in response to 3 yr of chronic N fertilization. The hardwood stand exhibited greater N limitation on biological function than the pine stand prior to fertilization as evidenced by a lower net N mineralization rate, nearly undetectable rates of net nitrification, and very low foliar N content. N additions were made in six equal applications throughout the growing season, and consisted of 5 and 15 g°m-2 °yr-1 of N as ammonium nitrate. The pine stand showed larger changes than the hardwood stand for extractable N, foliar N, nitrification, and N leaching loss. Retention of added N was essentially 100% for all but the high application pine plot from which significant N leaching occurred in the 3rd yr of application. From 75 to 92% of N added to fertilized plots was retained in the soil, with larger fractions retained in the hardwood stand than the pine stand for all treatments. As hypothesized, the stands are exhibiting highly nonlinear patterns of nitrogen output in response to continuous nitrogen inputs. The implications of this nonlinearity for regional eutrophication of surface waters and atmospheric deposition control policy are discussed.

FOREST BIOGEOCHEMISTRY AND PRIMARY PRODUCTION ALTERED BY NITROGEN SATURATION

Results from four intensive site-level manipulations and one extensive field survey in northern temperate and boreal forests show a consistent set of responses to chronic N additions. These include 1) initial and often large increase in net N mineralization followed by decreases, 2) increases in net nitrification. 3) increases in N concentration in foliage, and 4) decreased Mg∶N and Ca∶Al ratios, and declining tree growth and vigor in all evergreen stands. These results are synthesized into a set of proposed summary relationships that define the temporal pattern of responses of N-limited systems to N additions.

Long-term effects of experimental nitrogen additions on foliar litter decay and humus formation in forest ecosystems

Decomposition rates and N dynamics of foliar litter from 4 tree species were measured over a 72 month period on the Chronic Nitrogen Addition plots at the Harvard Forest, Petersham MA, beginning in November 1988. Plots received nitrogen additions of 0, 5 and 15 g N m-2yr-1 in two different stand types: red pine and mixed hardwood. Bags were collected in August and November of each year and litter analysed for mass remaining, nitrogen, cellulose and lignin content. Mass remaining was significantly greater for litter in nitrogen treated plots than in control plots after 48 months. Lignin content of litter was significantly higher with nitrogen treatments but there was little effect of treatment on cellulose content. N concentration was similar between treatments, but greater mass remaining in treated plots resulted in a higher total amount of N in humus produced in the high N plot. This mechanism could be a sink for up to 1.5 g N·m-2yr-1 of the 1.5 g N·m-2yr-1 added annually to the high N plots. Reduced decomposition rates in conjunction with increased lignin accumulation could impact global carbon sequestration as well.

The fate of 15N-labelled nitrate additions to a northern hardwood forest in eastern Maine, USA

We followed the movements of 15N-labelled nitrate additions into biomass and soil pools of experimental plots (15×15 m each) in a mid-successional beech-maple-birch-spruce forest in order to identify sinks for nitrate inputs to a forest ecosystem. Replicate plots (n=3) were spray-irrigated with either 28 or 56 kg N ha−1 year−1 using 15N-labelled nitric acid solutions (δ15N = 344‰ ) during four successive growing seasons (April–October). The 15N contents of foliage, bolewood, forests floor and mineral soil (0–5 cm) increased during the course of treatments. Mass balance calculations showed that one-fourth to one-third of the nitrate applied to forest plots was assimilated into and retained by above ground plant tissues and surface soil horizons at both rates of nitrate application. Plant and microbial assimilation were of approximately equal importance in retaining nitrate additions to this forest. Nitrate use among tree species varied, however, with red spruce showing lower rates of nitrate assimilation into foliage and bolewood than American beech and other deciduous species.

Consequences of climate change for biogeochemical cycling in forests of northeastern North America.

A critical component of assessing the impacts of climate change on forest ecosystems involves understanding associated changes in the biogeochemical cycling of elements. Evidence from research on northeastern North American forests shows that direct effects of climate change will evoke changes in biogeochemical cycling by altering plant physiology, forest productivity, and soil physical, chemical, and biological processes. Indirect effects, largely mediated by changes in species composition, length of growing season, and hydrology, will also be important. The case study presented here uses the quantitative biogeochemical model PnET-BGC to test assumptions about the direct and indirect effects of climate change on a northern hardwood forest ecosystem. Modeling results indicate an overall increase in net primary production due to a longer growing season, an increase in NO3– leaching due to large increases in net mineralization and nitrification, and slight declines in mineral weathering due to a reduction i...

Dissolved organic carbon and nitrogen relationships in forest litter as affected by nitrogen deposition.

Abstract Dissolved forms of carbon and nitrogen have become recognized for their importance in forest nutrient cycling. The role of dissolved organic carbon (DOC) as an energy source for microbial metabolism is of particular interest. A laboratory decomposition experiment was conducted to examine the relationship between potential increased N inputs (via acid deposition) and DOC production in the forest litter layer and subsequent effects on DOC availability in the forest floor. Air-dried leaf litter (seven species) was treated with nitrogen (nitrate or ammonium) or deionized (DI) water at weekly intervals throughout 15 weeks and leached with DI water at 1 or 2 week intervals. Leachate was analyzed for DOC, inorganic nitrogen (NO 3 − -N and NH 4 + -N) and dissolved organic nitrogen (DON). Litter was analyzed for percent C, percent N, weight loss and percent cellulose and lignin. Nitrogen treatments did not greatly affect DOC concentrations in litter leachate. Differences in DOC concentrations were primarily due to a wide range of initial litter chemistries, where species with high extractives and low lignin had the highest DOC leachate concentrations. Nitrogen treated samples showed greater weight loss than controls although nitrate and ammonium treatments were not significantly different. Between 6 and 39% of total carbon loss was leached as DOC. These findings suggest that different forest types could vary greatly in the quantity of carbon consumed or released and that nitrogen inputs appear to affect this overall cycle by increasing respiration (as measured by weight loss), rather than increasing DOC release into the soil solution. Further examination of the fate of DOC as it moves down in the soil profile and measurements of CO 2 evolution during laboratory decomposition, are necessary to better understand these processes.