Ecosystem Responses to Nitrogen Deposition in the Colorado Front Range

We asked whether 3–5 kg N y⁻¹ atmospheric N deposition was sufficient to have influenced natural, otherwise undisturbed, terrestrial and aquatic ecosystems of the Colorado Front Range by comparing ecosystem processes and properties east and west of the Continental Divide. The eastern side receives elevated N deposition from urban, agricultural, and industrial sources, compared with 1–2 kg N y⁻¹ on the western side. Foliage of east side old-growth Englemann spruce forests have significantly lower C:N and lignin:N ratios and greater N:Mg and N:P ratios. Soil % N is higher, and C:N ratios lower in the east side stands, and potential net N mineralization rates are greater. Lake NO₃ concentrations are significantly higher in eastern lakes than western lakes. Two east side lakes studied paleolimnologically revealed rapid changes in diatom community composition and increased biovolumes and cell concentrations. The diatom flora is now representative of increased disturbance or eutrophication. Sediment nitrogen isotopic ratios have become progressively lighter over the past 50 years, coincident with the change in algal flora, possibly from an influx of isotopically light N volatilized from agricultural fields and feedlots. Seventy-five percent of the increased east side soil N pool can be accounted for by increased N deposition commensurate with human settlement. Nitrogen emissions from fixed, mobile, and agricultural sources have increased dramatically since approximately 1950 to the east of the Colorado Front Range, as they have in many parts of the world. Our findings indicate even slight increases in atmospheric deposition lead to measurable changes in ecosystem properties. Received 16 November 1999; accepted 8 February 2000.     

Large Loss of Dissolved Organic Nitrogen from Nitrogen-Saturated Forests in Subtropical China

Dissolved organic nitrogen (DON) has recently been recognized as an important component of terrestrial N cycling, especially under N-limited conditions; however, the effect of increased atmospheric N deposition on DON production and loss from forest soils remains controversial. Here we report DON and dissolved organic carbon (DOC) losses from forest soils receiving very high long-term ambient atmospheric N deposition with or without additional experimental N inputs, to investigate DON biogeochemistry under N-saturated conditions. We studied an old-growth forest, a young pine forest, and a young mixed pine/broadleaf forest in subtropical southern China. All three forests have previously been shown to have high nitrate (NO₃⁻) leaching losses, with the highest loss found in the old-growth forest. We hypothesized that DON leaching loss would be forest specific and that the strongest response to experimental N input would be in the N-saturated old-growth forest. Our results showed that under ambient deposition (35–50 kg N ha⁻¹ y⁻¹ as throughfall input), DON leaching below the major rooting zone in all three forests was high (6.5–16.9 kg N ha⁻¹ y⁻¹). DON leaching increased 35–162% following 2.5 years of experimental input of 50–150 kg N ha⁻¹ y⁻¹. The fertilizer-driven increase of DON leaching comprised 4–17% of the added N. A concurrent increase in DOC loss was observed only in the pine forest, even though DOC:DON ratios declined in all three forests. Our data showed that DON accounted for 23–38% of total dissolved N in leaching, highlighting that DON could be a significant pathway of N loss from forests moving toward N saturation. The most pronounced N treatment effect on DON fluxes was not found in the old-growth forest that had the highest DON loss under ambient conditions. DON leaching was highly correlated with NO₃⁻ leaching in all three forests. We hypothesize that abiotic incorporation of excess NO₃⁻ (through chemically reactive NO₂⁻) into soil organic matter and the consequent production of N-enriched dissolved organic matter is a major mechanism for the consistent and large DON loss in the N-saturated subtropical forests of southern China. Dr. YT Fang performed research, analyzed data, and wrote the paper; Prof. WX Zhu participated in the initial experimental design, analyzed data, and took part in writing the paper; Prof. P Gundersen conceived the study and took part in writing; Prof. JM Mo and Prof. GY Zhou conceived study; Prof. M Yoh analyzed part of the data and contributed to the development of DON model.     

Symptoms of nitrogen saturation in an aggrading forested watershed in western Maryland

The objective of this study was to evaluate the nitrogen (N) biogeochemistry of an 18–22 year old forested watershed in western Maryland. We hypothesized that this watershed should not exhibit symptoms of N saturation. This watershed was a strong source of nitrate (NO₃ ⁻) to the stream in all years, with a mean annual export of 9.5 kg N ha⁻¹ year⁻¹ and a range of 4.4–18.4 kg N ha⁻¹ year⁻¹. During the 2001 and 2002 water years, wet deposition of inorganic N was 9.0 kg N ha⁻¹ year⁻¹ and 6.3 kg N ha⁻¹ year⁻¹, respectively. Watershed N export rates in 2001 and 2002 water years were 4.2 kg N ha⁻¹ year⁻¹ and 5.3 kg N ha⁻¹ year⁻¹, respectively. During the wetter water years of 2003 and 2004, the watershed exported 15.0 kg N ha⁻¹ year⁻¹ and 18.4 kg N ha⁻¹ year⁻¹, rates that exceeded annual wet deposition of N by a factor of two (7.5 kg N ha⁻¹ year⁻¹ in 2003) and three (5.5 kg N ha⁻¹ year⁻¹ in 2004). Consistent with the high rates of N export, were high concentrations (2.1–3.3%) of N in foliage, wood (0.3%) and fine roots, low C:N ratios in the forest floor (17–24) and mineral soil (14), high percentages (83–96%) of the amount of mineralized N that was nitrified and elevated N concentrations (up to 3 mg N l⁻¹) in soil solution. Although this watershed contained a young aggrading forest, it exhibited several symptoms of N saturation commonly observed in more mature forests.     

The Long-term Effects of Disturbance on Organic and Inorganic Nitrogen Export in the White Mountains, New Hampshire

Traditional biogeochemical theories suggest that ecosystem nitrogen retention is controlled by biotic N limitation, that stream N losses should increase with successional age, and that increasing N deposition will accelerate this process. These theories ignore the role of dissolved organic nitrogen (DON) as a mechanism of N loss. We examined patterns of organic and inorganic N export from sets of old-growth and historically (80–110 years ago) logged and burned watersheds in the northeastern US, a region of moderate, elevated N deposition. Stream nitrate concentrations were strongly seasonal, and mean (± SD) nitrate export from old-growth watersheds (1.4 ± 0.6 kg N ha⁻¹ y⁻¹) was four times greater than from disturbed watersheds (0.3 ± 0.3 kg N ha⁻¹ y⁻¹), suggesting that biotic control over nitrate loss can persist for a century. DON loss averaged 0.7 (± 0.2) kg N ha⁻¹ y⁻¹ and accounted for 28–87% of total dissolved N (TDN) export. DON concentrations did not vary seasonally or with successional status, but correlated with dissolved organic carbon (DOC), which varied inversely with hardwood forest cover. The patterns of DON loss did not follow expected differences in biotic N demand but instead were consistent with expected differences in DOC production and sorption. Despite decades of moderate N deposition, TDN export was low, and even old-growth forests retained at least 65% of N inputs. The reasons for this high N retention are unclear: if due to a large capacity for N storage or biological removal, N saturation may require several decades to occur; if due to interannual climate variability, large losses of nitrate may occur much sooner. Received 27 April 1999; accepted 30 May 2000.     

Cloudwater Inputs of Nitrogen to Forest Ecosystems in Southern Chile: Forms, Fluxes, and Sources

Nitrogen (N) has been considered a limiting nutrient to many aquatic and terrestrial ecosystems. However, human activity has resulted in increased atmospheric N deposition worldwide such that N pollution is now altering ecosystem function in many locations. Research on atmospheric deposition has focused primarily on inorganic nitrogen (DIN; NH₄ ⁺-N + NO₃ ⁻-N) via rainwater and dry deposition as the main N input to ecosystems. Recently, organic N (ON) has been shown to be an important constituent in rainwater or dry deposition. Here we show that ON dominated (66%) total N in cloudwater from a remote site in southern Chile. Cloudwater from more human-impacted sites in northeastern USA had lower ON concentrations and DIN was dominant. We estimate that cloudwater delivers up to 2 kg ha⁻¹ DIN and 9 kg ha⁻¹ ON annually, compared to less than 1 kg ha⁻¹ of DIN deposition via rainwater, thus it may contribute substantially to the N-economy of Chilean coastal forests. We also suggest that the adjacent ocean, where biologic productivity is high, may be a major source of N in Chilean cloudwater. This proposed marine-terrestrial flux via cloud deposition has not previously been identified and may be an example of the ocean feeding the forest. We suggest that this type of cross boundary flux may be common in other upwelling zones, such as along the west coasts of Africa, North and South America and east India, and warrants further substantiation and investigation. Received 30 June 2000; accepted 18 October 2000     

Nitrogen uptake in a Norway spruce stand following ammonium sulphate application,fertigation, irrigation,drought and nitrogen-free-fertilisation

The above-ground accumulation of N,N uptake and litter quality resulting from improved or deteriorated availability of water and nutrients in a 25 year old Norway spruce stand in SW Sweden (as part of the Skogaby project) is presented. Treatment include irrigation; artificial drought; ammonium sulphate addition; N-free-fertilisation and irrigation with liquid fertilisers including a complete set of nutrients according to the Ingested principle (fertigation). At start of the experiment the stand contained 86.5 t dry mass and 352 kg N ha⁻¹. The following three years the annual N uptake in untreated trees was 32 kg N ha⁻¹ to be compared with the annual N throughfall of 17 kg ha⁻¹. Simultaneously, the treatment with ammonium sulphate and liquid fertilisation resulted in 48 and 56 kg ha⁻¹ y⁻¹, respectively, in treatment specific N-uptake following an application of 100 kg N ha⁻¹ y⁻¹. Addition of a N-free fertiliser resulted in improved N-uptake by 19 kg N ha⁻¹ y⁻¹ and irrigation by 10 kg N ha⁻¹ y⁻¹, compared to control. A linear relation between total above-ground dry mass production and N-uptake was found for trees growing with similar water availability. Dry mass production increased with increased water availability given the same N-uptake. It is concluded that the studied stand this far is not N saturated', as N fertilisation resulted in both increased N uptake and increased growth. Addition of a N-free-fertiliser resulted in increased uptake of N compared to the control, indicating an increased mineralisation rate or uptake capacity of the root system. The linear relation between N uptake and biomass production shows that at this study site N is a highly limiting factor for growth.     

Response of Litter Decomposition to Simulated N Deposition in Disturbed, Rehabilitated and Mature Forests in Subtropical China

The response of decomposition of litter for the dominant tree species in disturbed (pine), rehabilitated (pine and broadleaf mixed) and mature (monsoon evergreen broadleaf) forests in subtropical China to simulated N deposition was studied to address the following hypothesis: (1) litter decomposition is faster in mature forest (high soil N availability) than in rehabilitated/disturbed forests (low soil N availability); (2) litter decomposition is stimulated by N addition in rehabilitated and disturbed forests due to their low soil N availability; (3) N addition has little effect on litter decomposition in mature forest due to its high soil N availability. The litterbag method (a total of 2880 litterbags) and N treatments: Control-no N addition, Low-N: −5 g N m⁻² y⁻¹, Medium-N: −10 g N m⁻² y⁻¹, and High-N: −15 g N m⁻² y⁻¹, were employed to evaluate decomposition_. Results indicated that mature forest, which has likely been N saturated due to both long-term high N deposition in the region and the age of the ecosystem, had the highest litter decomposition rate, and exhibited no significant positive and even some negative response to nitrogen additions. However, both disturbed and rehabilitated forests, which are still N limited due to previous land use history, exhibited slower litter decomposition rates with significant positive effects from nitrogen additions. These results suggest that litter decomposition and its responses to N addition in subtropical forests of China vary depending on the nitrogen status of the ecosystem.     

Nitrogen Fixation by Termites in Tropical Forests, Thailand

Nitrogen (N) fixed by termites was evaluated as a N input to decomposition processes in two tropical forests, a dry deciduous forest (DDF) and the neighboring dry evergreen forest (DEF), Thailand. A diverse group of termite species were assayed by acetylene reduction method and only the wood/litter-feeding termites were found to fix N. More intensive samplings of two abundant species, Microcerotermes crassus and Globitermes sulphureus, were done across several seasons, suggesting N fixation rates of 0.21 and 0.28 kg ha⁻¹ y⁻¹ by termites in the DDF and DEF, respectively. Also, estimates of asymbiotic N fixation rates were 0.75 and 3.95 kg ha⁻¹ y⁻¹. N fixed by termites and by asymbiotic fixers is directly supplied to decomposers breaking down dead plant material and could be a major source of their N. N fixed by termites was 7–22% of that fixed by termites and asymbiotic fixers. Although N fixed by termites is a small input compared to other inputs, this N is likely important for decomposition processes.     

Soil Inorganic N Leaching in Edges of Different Forest Types Subject to High N Deposition Loads

We report on soil leaching of dissolved inorganic nitrogen (DIN) along transects across exposed edges of four coniferous and four deciduous forest stands. In a 64-m edge zone, DIN leaching below the main rooting zone was enhanced relative to the interior (at 128 m from the edge) by 21 and 14 kg N ha⁻¹ y⁻¹ in the coniferous and deciduous forest stands, respectively. However, the patterns of DIN leaching did not univocally reflect those of DIN throughfall deposition. DIN leaching in the first 20 m of the edges was lower than at 32–64 m from the edge (17 vs. 36 kg N ha⁻¹ y⁻¹ and 15 vs. 24 kg N ha⁻¹ y⁻¹ in the coniferous and deciduous forests, respectively). Nitrogen stocks in the mineral topsoil (0–30 cm) were, on average, 943 kg N ha⁻¹ higher at the outer edges than in the interior, indicating that N retention in the soil is probably one of the processes involved in the relatively low DIN leaching in the outer edges. We suggest that a complex of edge effects on biogeochemical processes occurs at the forest edges as a result of the interaction between microclimate, tree dynamics (growth and litterfall), and atmospheric deposition of N and base cations.     

Long-Term Trends in Stream Nitrate Concentrations and Losses Across Watersheds Undergoing Recovery from Acidification in the Czech Republic

Stream nitrogen (N) export and nitrate concentration were measured at 14 forested watersheds (GEOMON network) in the Czech Republic between 1994 and 2005. In the last several decades, emissions of sulfur (S) and N compounds have decreased throughout much of Europe. In the Czech Republic, atmospheric deposition of S has decreased substantially since the beginning of 1990s, whereas N deposition remains largely unchanged at most sites. The mean dissolved inorganic nitrogen (DIN) streamwater export ranged from 0.2 to 12.2 kg ha⁻¹ y⁻¹ at the GEOMON sites. Despite decades of elevated N deposition, 44–98% of DIN inputs to these watersheds were retained or denitrified, and many watersheds showed seasonal variation in nitrate concentrations. Dissolved organic N export was quantified in 1 year only and ranged from 0.05 to 3.5 kg ha⁻¹ y⁻¹. Spatial variability in DIN export among watersheds was best explained by spatial variability in average acidic deposition, particularly S deposition (R ² = 0.81, P < 0.001); DIN input and forest floor carbon:nitrogen (C/N) also provided significant explanatory power. DIN export was strongly influenced by the forest floor C/N ratio and depth of the forest floor soils (R ² = 0.72, P < 0.001). The only variable that predicted variations in forest floor C/N (R ² = 0.32, P < 0.05) among watersheds was S deposition. Forest floor depth was also related to deposition variables, with S deposition providing the most explanatory power (R ² = 0.50, P < 0.01). Variation in forest floor depth was also associated with climatic factors (precipitation and temperature). Temporal variability in DIN export was primarily associated with changes in acidic deposition over time; S deposition explained 41% of variability in DIN exports among all watersheds and years. Extensive acidification of forested watersheds was associated with the extraordinarily high S inputs to much of the Czech Republic during earlier decades. We hypothesize that recovery from acidification has led to improved tree health as well as enhanced microbial activity in the forest floor. As these watersheds move into a new regime with dramatically lower sulfur inputs, we expect continued declines in nitrate output.