Fine root turnover in forest ecosystems in relation to quantity and form of nitrogen availability: a comparison of two methods

Summary Two methods of estimating fine root production and turnover are compared for 13 forest ecosystems exhibiting a wide range in form (NH₄ ⁺ vs. NO₃ ^-) and quantity of available nitrogen. The two methods are by comparison of seasonal maxima and minima in biomess and by nitrogen budgeting. Both methods give similar results for stands with low rates of nitrification. The budgeting method predicts higher fine root turnover and productivity than the max-min method for systems with significant rates of nitrification.     

Sources of nitrate in rivers draining sixteen watersheds in the northeastern U.S.: Isotopic constraints

The feasibility of using nitrogen and oxygenisotope ratios of nitrate (NO₃ ^–) forelucidating sources and transformations ofriverine nitrate was evaluated in a comparativestudy of 16 watersheds in the northeastern U.S.A. Stream water was sampled repeatedly at theoutlets of the watersheds between January andDecember 1999 for determining concentrations,¹⁵N values, and ¹⁸Ovalues of riverine nitrate.In conjunction with information about land useand nitrogen fluxes,¹⁵N_nitrate and¹⁸O_nitrate values providedmainly information about sources of riverinenitrate. In predominantly forested watersheds,riverine nitrate had mean concentrations ofless than 0.4 mg NO₃ ^–-N L^–1,¹⁵N_nitrate values of lessthan +5, and ¹⁸O_nitratevalues between +12 and +19. This indicatesthat riverine nitrate was almost exclusivelyderived from soil nitrification processes withpotentially minor nitrate contributions fromatmospheric deposition in some catchments. Inwatersheds with significant agricultural andurban land use, concentrations of riverinenitrate were as high as 2.6 mg NO₃ ^–-NL^–1 with ¹⁵N_nitratevalues between +5 and +8 and¹⁸O_nitrate values generallybelow +15. Correlations between nitrateconcentrations, ¹⁵N_nitratevalues, and N fluxes suggest that nitrate inwaste water constituted a major, and nitrate inmanure a minor additional source of riverinenitrate. Atmospheric nitrate deposition ornitrate-containing fertilizers were not asignificant source of riverine nitrate inwatersheds with significant agricultural andurban land use. Although complementary studiesindicate that in-stream denitrification wassignificant in all rivers, the isotopiccomposition of riverine nitrate sampled at theoutlet of the 16 watersheds did not provideevidence for denitrification in the form ofelevated ¹⁵N_nitrate and¹⁸O_nitrate values. Relativelylow isotopic enrichment factors for nitrogenand oxygen during in-stream denitrification andcontinuous admixture of nitrate from theabove-described sources are thought to beresponsible for this finding.     

Natural<Superscript>15</Superscript> N Abundance of Plants and Soil N in a Temperate Coniferous Forest

Measurement of nitrogen isotopic composition (¹⁵N) of plants and soil nitrogen might allow the characteristics of N transformation in an ecosystem to be detected. We tested the measurement of ¹⁵N for its ability to provide a picture of N dynamics at the ecosystem level by doing a simple comparison of ¹⁵N between soil N pools and plants, and by using an existing model. ¹⁵N of plants and soil N was measured together with foliar nitrate reductase activity (NRA) and the foliar NO₃^– pool at two sites with different nitrification rates in a temperature forest in Japan. ¹⁵N of plants was similar to that of soil NO₃^– in the high-nitrification site. Because of high foliar NRA and the large foliar NO₃^– pool at this site, we concluded that plant ¹⁵N indicated a great reliance of plants on soil NO₃^– there. However, many ¹⁵N of soil N overlapped each other at the other site, and ¹⁵N could not provide definitive evidence of the N source. The existing model was verified by measured ¹⁵N of soil inorganic N and it explained the variations of plant ¹⁵N between the two sites in the context of relative importance of nitrification, but more information about isotopic fractionations during plant N uptake is required for quantitative discussions about the plant N source. The model applied here can provide a basis to compare ¹⁵N signatures from different ecosystems and to understand N dynamics.     

Climate and species affect fine root production with long-term fertilization in acidic tussock tundra near Toolik Lake,Alaska

Long-term fertilization of acidic tussock tundra has led to changes in plant species composition, increases in aboveground production and biomass and substantial losses of soil organic carbon (SOC). Root litter is an important input to SOC pools, although little is known about fine root demography in tussock tundra. In this study, we examined the response of fine root production and live standing fine root biomass to short- and long-term fertilization, as changes in fine root demography may contribute to observed declines in SOC. Live standing fine root biomass increased with long-term fertilization, while fine root production declined, reflecting replacement of the annual fine root system of Eriophorum vaginatum, with the long-lived fine roots of Betula nana. Fine root production increased in fertilized plots during an unusually warm growing season, but remained unchanged in control plots, consistent with observations that B. nana shows a positive response to climate warming. Calculations based on a few simple assumptions suggest changes in fine root demography with long-term fertilization and species replacement could account for between 20 and 39% of the observed declines in SOC stocks.     

Forest nitrogen sinks in large eastern U.S. watersheds: estimates from forest inventory and an ecosystem model

The eastern U.S. receives elevated rates of Ndeposition compared to preindustrial times, yetrelatively little of this N is exported indrainage waters. Net uptake of N into forestbiomass and soils could account for asubstantial portion of the difference between Ndeposition and solution exports. We quantifiedforest N sinks in biomass accumulation andharvest export for 16 large river basins in theeastern U.S. with two separate approaches: (1)using growth data from the USDA ForestService's Forest Inventory and Analysis (FIA)program, and (2) using a model of forestnitrogen cycling (PnET-CN) linked to FIAinformation on forest age-class structure. Themodel was also used to quantify N sinks in soiland dead wood, and nitrate losses below therooting zone. Both methods agreed that netgrowth rates were highest in the relativelyyoung forests on the Schuylkill watershed, andlowest in the cool forests of northern Maine. Across the 16 watersheds, wood export removedan average of 2.7 kg N ha^–1 yr^–1(range: 1–5 kg N ha^–1 yr^–1), andstanding stocks increased by 4.0 kg N ha^–1yr^–1 (–3 to 8 kg N ha^–1 yr^–1). Together, these sinks for N in woody biomassamounted to a mean of 6.7 kg N ha^–1yr^–1 (2–9 kg N ha^–1 yr^–1), or73% (15–115%) of atmospheric N deposition. Modeled rates of net N sinks in dead wood andsoil were small; soils were only a significantnet sink for N during simulations ofreforestation of degraded agricultural sites. Predicted losses of nitrate depended on thecombined effects of N deposition, and bothshort- and long-term effects of disturbance. Linking the model with forest inventoryinformation on age-class structure provided auseful step toward incorporating realisticpatterns of forest disturbance status acrossthe landscape.     

Chronic nitrogen additions suppress decomposition and sequester soil carbon in temperate forests

The terrestrial biosphere sequesters up to a third of annual anthropogenic carbon dioxide emissions, offsetting a substantial portion of greenhouse gas forcing of the climate system. Although a number of factors are responsible for this terrestrial carbon sink, atmospheric nitrogen deposition contributes by enhancing tree productivity and promoting carbon storage in tree biomass. Forest soils also represent an important, but understudied carbon sink. Here, we examine the contribution of trees versus soil to total ecosystem carbon storage in a temperate forest and investigate the mechanisms by which soils accumulate carbon in response to two decades of elevated nitrogen inputs. We find that nitrogen-induced soil carbon accumulation is of equal or greater magnitude to carbon stored in trees, with the degree of response being dependent on stand type (hardwood versus pine) and level of N addition. Nitrogen enrichment resulted in a shift in organic matter chemistry and the microbial community such that unfertilized soils had a higher relative abundance of fungi and lipid, phenolic, and N-bearing compounds; whereas, N-amended plots were associated with reduced fungal biomass and activity and higher rates of lignin accumulation. We conclude that soil carbon accumulation in response to N enrichment was largely due to a suppression of organic matter decomposition rather than enhanced carbon inputs to soil via litter fall and root production.     

Molecular-level changes in soil organic matter composition after 10 years of litter,root and nitrogen manipulation in a temperate forest

With climate change, forests are expected to receive increased inputs of carbon (C) and nitrogen (N) but it is unclear how this will modify forest C cycling and storage at the molecular-level. To investigate the response of forest soil organic matter (SOM) to changes in soil inputs, a study area was established in a Michigan hardwood forest as part of the Detrital Input and Removal Treatments (DIRT) network. Experimental treatments were comprised of both exclusions of detrital inputs (No Litter, No Roots, No Inputs) and additions of C and N (Double Litter, N-Addition, Double Litter?+?N, Wood). After 10 years of treatment, the soils were characterized using elemental analysis, molecular biomarker techniques, nuclear magnetic resonance spectroscopy, and microbial biomass C measurements. Although manipulation of detrital inputs did not significantly change the soil C and N content after 10 years, alterations in the cycling and distribution of SOM components were observed. Root exclusion enhanced SOM degradation, while doubling litter favoured the degradation of more labile forms of soil C such as unsaturated n-alkanoic acids and simple sugars. N-Addition and Double Litter?+?N increased the concentrations of extractable biomarkers, including aliphatic and cyclic lipids and compounds derived from cutin, suberin, and lignin. Microbial biomass C also varied with experimental litter input manipulations and N addition, and these data were consistent with the observed changes in SOM composition. Overall, the observed shifts in SOM chemistry after 10 years of manipulating ecosystem inputs highlight the sensitivity of natural systems to changes in amounts of C and N inputs from roots and litter, and N inputs from external sources.     

Immobilization of a 15N-labeled nitrate addition by decomposing forest litter

This paper explores the biological consequences of climate change by integrating the results of a tripartite investigation involving fumarole, field manipulation and laboratory incubation experiments. The geographical region for this research is the maritime Antarctic. Under contemporary climate conditions, the lithosols in this region support only a sparse cryptogamic flora of limited taxonomic diversity and low structural complexity. However, the existence in geothermal areas of temperate species (e.g. Campylopus introflexus, Marchantia polymorpha, Philonotis acicularis) growing outside their normal biogeographical range suggests that elevated temperature and humidity may alter the trajectory of community development towards Magellanic or Patagonian composition. Productivity is also likely to increase, as indicated by significantly greater vegetative biomass recorded beneath climate-ameliorating soil covers than in controls. Barren fellfield soil samples transplanted to the laboratory and incubated at temperatures of 2–25°C show rapid development of moss, algae and lichen propagules in the range 15–25°C. A variety of species develop that have not been recorded in the field. The presence of exotic taxa indicates the existence of a dormant propagule bank in maritime Antarctic soils and suggests that no significant delay is likely to occur between the onset of climate warming and community development: instead, rapid establishment of those species favoured by the new climate conditions will yield a distinct founder effect, with increasing above- and below-ground biomass stimulating biogeochemical cycling. It is argued that the combined results of this synthesis identify generic responses to climate change arising from the importance at high latitudes of low temperature and water availability as limiting factors: subject to other growth resources being non-limiting, a more consistent stimulatory response to climate change may be expected than in temperate or tropical regions. The tripartite approach, encompassing field, microcosm and laboratory methodologies, renders the conclusions more robust than any single study considered in isolation.     

15N natural abundances and N use by tundra plants

Plant species collected from tundra ecosystems located along a north-south transect from central Alaska to the north coast of Alaska showed large and consistent differences in ¹⁵N natural abundances. Foliar ¹⁵N values varied by about 10% among species within each of two moist tussock tundra sites. Differences in ¹⁵N contents among species or plant groups were consistent across moist tussock tundra at several other sites and across five other tundra types at a single site. Ericaceous species had the lowest ¹⁵N values, ranging between about –8 to –6. Foliar ¹⁵N contents increased progressively in birch, willows and sedges to maximum ¹⁵N values of about +2 in sedges. Soil ¹⁵N contents in tundra ecosystems at our two most intensively studied sites increased with depth and ¹⁵N values were usually higher for soils than for plants. Isotopic fractionations during soil N transformations and possibly during plant N uptake could lead to observed differences in ¹⁵N contents among plant species and between plants and soils. Patterns of variation in ¹⁵N content among species indicate that tundra plants acquire nitrogen in extremely nutrient-poor environments by competitive partitioning of the overall N pool. Differences in plant N sources, rooting depth, mycorrhizal associations, forms of N taken up, and other factors controlling plant N uptake are possible causes of variations in ¹⁵N values of tundra plant species.     

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 (¹⁵N = 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.