Nitrogen yields from undisturbed watersheds in the Americas

Yields of total fixed nitrogen and nitrogen fractions are summarized for thirty-one watersheds in which anthropogenic disturbance of the nitrogen cycle, either through land use or atmospheric deposition, is negligible or slight. These yields are taken as representative of background conditions over a broad range of watershed areas, elevations, and vegetation types. The data set focuses on watersheds of the American tropics, but also includes information on the Gambia River (Africa) and some small watersheds in the Sierra Nevada of California. For the tropical watersheds, total nitrogen yield averages 5.1 kg ha^–1 y^–1. On average, 30% of the total is particulate and 70% is dissolved. Of the dissolved fraction, an average of 50% is organic and 50% is inorganic, of which 20% is ammonium and 80% is nitrate. Yields are substantially lower than previously estimated for background conditions. Yields of all nitrogen fractions are strongly related to runoff, which also explains a large percentage of variance in yield of total nitrogen (r²=0.85). For total nitrogen and nitrogen fractions, yield increases at about two-thirds the rate of runoff; concentration decreases as runoff increases. There is a secondary but significant positive relationship between elevation and yield of DIN. Ratios DON/TDN and PN/TN both are related to watershed area rather than runoff; DON/TDN decreases and PN/TN increases toward higher stream orders. The analysis suggests for tropical watersheds the existence of mechanisms promoting strong homeostasis in the yield of N and its fractions for a given moisture regime, as well as predictable downstream change in proportionate representation N fractions. Yields and concentrations for small tropical watersheds are much larger than for the few temperate ones with which comparisons are possible.     

Riparian nitrogen dynamics in two geomorphologically distinct tropical rain forest watersheds: subsurface solute patterns

Nitrate, ammonium, dissolved organic N, and dissolved oxygen were measured in stream water and shallow groundwater in the riparian zones of two tropical watersheds with different soils and geomorphology. At both sites, concentrations of dissolved inorganic N (DIN; NH₄ ⁺- and NO₃ ⁻-N) were low in stream water (< 110 ug/L). Markedly different patterns in DIN were observed in groundwater collected at the two sites. At the first site (Icacos watershed), DIN in upslope groundwater was dominated by NO₃ ⁻-N (550 ug/L) and oxygen concentrations were high (5.2 mg/L). As groundwater moved through the floodplain and to the stream, DIN shifted to dominance by NH₄ ⁺-N (200–700 ug/L) and groundwater was often anoxic. At the second site (Bisley watershed), average concentrations of total dissolved nitrogen were considerably lower (300 ug/L) than at Icacos (600 ug/L), and the dominant form of nitrogen was DON rather than inorganic N. Concentrations of NH₄ ⁺ and NO₃ ⁻ were similar throughout the riparian zone at Bisley, but concentrations of DON declined from upslope wells to stream water. Differences in speciation and concentration of nitrogen in groundwater collected at the two sites appear to be controlled by differences in redox conditions and accessibility of dissolved N to plant roots, which are themselves the result of geomorphological differences between the two watersheds. At the Icacos site, a deep layer of coarse sand conducts subsurface water to the stream below the rooting zone of riparian vegetation and through zones of strong horizontal redox zonation. At the Bisley site, infiltration is impeded by dense clays and saturated flow passes through the variably oxidized rooting zone. At both sites, hydrologic export of nitrogen is controlled by intense biotic activity in the riparian zone. However, geomorphology appears to strongly modify the importance of specific biotic components.     

The influence of climate on average nitrogen export from large watersheds in the Northeastern United States

The flux of nitrogen in large rivers in North America and Europe is well explained as a function of the net anthropogenic inputs of nitrogen to the landscape, with on average 20 to 25% of these inputs exported in rivers and 75 to 80% of the nitrogen retained or denitrified in the landscape. Here, we use data for average riverine nitrogen fluxes and anthropogenic inputs of nitrogen over a 6-year period (1988–1993) for 16 major watersheds in the northeastern United States to examine if there is also a climatic influence on nitrogen fluxes in rivers. Previous studies have shown that for any given river, nitrogen fluxes are greater in years with higher discharge, but this can be interpreted as storage of nitrogen in the landscape during dry years and flushing of this stored nitrogen during wet years. Our analyses demonstrate that there is also a longer-term steady-state influence of climate on riverine nitrogen fluxes. Those watersheds that have higher precipitation and higher discharge export a greater fraction of the net anthropogenic inputs of nitrogen. This fractional export ranges from 10 to 15% of the nitrogen inputs in drier watersheds in the northeastern United States to over 35% in the wetter watersheds. We believe this is driven by lower rates of denitrification in the wetter watersheds, perhaps because shorter water residence times do not allow for as much denitrification in riparian wetlands and low-order streams. Using mean projections for the consequences of future climate change on precipitation and discharge, we estimate that nitrogen fluxes in the Susquehanna River to Chesapeake Bay may increase by 3 to 17% by 2030 and by 16 to 65% by 2095 due to greater fractional delivery of net anthropogenic nitrogen inputs as precipitation and discharge increase. Although these projections are highly uncertain, they suggest a need to better consider the influence of climate on riverine nitrogen fluxes as part of management efforts to control coastal nitrogen pollution.     

A test of a potential short cut in the nitrogen cycle: The role of exudation of symbiotically fixed nitrogen from the roots of a N-fixing tree and the effects of increased atmospheric CO2 and temperature

N-fixing trees facilitate the growth of neighboring trees of other species. These neighboring species benefit from the simple presence of the N fixation symbiosis in their surroundings. Because of this phenomenon, it has been hypothesized that a change in atmospheric CO₂ concentration may alter the role of N-fixing trees in their environment. It is thought that the role of N-fixing trees in ecosystems of the future may be more important since they may help sustain growth increases due to increased CO₂ concentration in nitrogen limited forests. We examined: (1) whether symbiotically fixed N was exuded from roots, (2) whether a doubled atmospheric CO₂ concentration would result in increased organic N exudation from roots, and (3) whether increased temperature or N availability affected N exudation from roots. This study analyzed exudation of dissolved organic N from the roots of seedlings of the N-fixing tree Robinia pseudoacacia L. in a full factorial design with 2 CO₂ (35.0 and 70.0 Pa) × 2 temperature (26 or 30 °C during the day) × 2 N fertilizer (0 and 10.0 mM N concentration) levels. Trees with no other source of N except N fixation exuded about 1% to 2% of the fixed N through their roots as dissolved organic N. Increased atmospheric CO₂ concentrations did not, however, increase N exudation rates on a per gram belowground biomass basis. A 4 °C increase in temperature and N fertilization did, however, significantly increase N exudation rates. These results suggest that exudation of dissolved organic N from roots or nodules of N-fixing trees could be a significant, but minor, pathway of transferring N to neighboring plants in a much more rapid and direct way than cycling through death, decomposition and mineralization of plant residues. And, while exudation rates of dissolved organic N from roots were not significantly affected by atmospheric CO₂ concentration, the previously observed CO₂ fertilization effect on N-fixing trees suggests that N exudation from roots could play a significant but minor role in sustaining increases in forest growth, and thus C storage, in a CO₂ enriched atmosphere.     

Effects of Succession on Nitrogen Export in the West-Central Cascades, Oregon

This study examined impacts of succession on N export from 20 headwater stream systems in the west central Cascades of Oregon, a region of low anthropogenic N inputs. The seasonal and successional patterns of nitrate (NO₃−N) concentrations drove differences in total dissolved N concentrations because ammonium (NH₄−N) concentrations were very low (usually < 0.005 mg L⁻¹) and mean dissolved organic nitrogen (DON) concentrations were less variable than nitrate concentrations. In contrast to studies suggesting that DON levels strongly dominate in pristine watersheds, DON accounted for 24, 52, and 51% of the overall mean TDN concentration of our young (defined as predominantly in stand initiation and stem exclusion phases), middle-aged (defined as mixes of mostly understory reinitiation and older phases) and old-growth watersheds, respectively. Although other studies of cutting in unpolluted forests have suggested a harvest effect lasting 5 years or less, our young successional watersheds that were all older than 10 years still lost significantly more N, primarily as NO₃−N, than did watersheds containing more mature forests, even though all forest floor and mineral soil C:N ratios were well above levels reported in the literature for leaching of dissolved inorganic nitrogen. The influence of alder may contribute to these patterns, although hardwood cover was quite low in all watersheds; it is possible that in forested ecosystems with very low anthropogenic N inputs, even very low alder cover in riparian zones can cause elevated N exports. Only the youngest watersheds, with the highest nitrate losses, exhibited seasonal patterns of increased summer uptake by vegetation as well as flushing at the onset of fall freshets. Older watersheds with lower N losses did not exhibit seasonal patterns for any N species. The results, taken together, suggest a role for both vegetation and hydrology in N retention and loss, and add to our understanding of N cycling by successional forest ecosystems influenced by disturbance at various spatial and temporal scales in a region of relatively low anthropogenic N input.     

Benthic organic carbon influences denitrification in streams with high nitrate concentration

1. Anthropogenic activities have increased reactive nitrogen availability, and now many streams carry large nitrate loads to coastal ecosystems. Denitrification is potentially an important nitrogen sink, but few studies have investigated the influence of benthic organic carbon on denitrification in nitrate‐rich streams. 2. Using the acetylene‐block assay, we measured denitrification rates associated with benthic substrata having different proportions of organic matter in agricultural streams in two states in the mid‐west of the U.S.A., Illinois and Michigan. 3. In Illinois, benthic organic matter varied little between seasons (5.9–7.0% of stream sediment), but nitrate concentrations were high in summer (>10 mg N L⁻¹) and low (<0.5 mg N L⁻¹) in autumn. Across all seasons and streams, the rate of denitrification ranged from 0.01 to 4.77 μg N g⁻¹ DM h⁻¹ and was positively related to stream‐water nitrate concentration. Within each stream, denitrification was positively related to benthic organic matter only when nitrate concentration exceeded published half‐saturation constants. 4. In Michigan, streams had high nitrate concentrations and diverse benthic substrata which varied from 0.7 to 72.7% organic matter. Denitrification rate ranged from 0.12 to 11.06 μg N g⁻¹ DM h⁻¹ and was positively related to the proportion of organic matter in each substratum. 5. Taken together, these results indicate that benthic organic carbon may play an important role in stream nitrogen cycling by stimulating denitrification when nitrate concentrations are high.     

Dissolved organic nitrogen budgets for upland, forested ecosystems in New England

Relatively high deposition ofnitrogen (N) in the northeastern United States hascaused concern because sites could become N saturated.In the past, mass-balance studies have been used tomonitor the N status of sites and to investigate theimpact of increased N deposition. Typically, theseefforts have focused on dissolved inorganic forms ofN (DIN = NH₄-N + NO₃-N) and have largelyignored dissolved organic nitrogen (DON) due todifficulties in its analysis. Recent advances in themeasurement of total dissolved nitrogen (TDN) havefacilitated measurement of DON as the residual of TDN– DIN. We calculated DON and DIN budgets using data onprecipitation and streamwater chemistry collected from9 forested watersheds at 4 sites in New England. TDNin precipitation was composed primarily of DIN. Netretention of TDN ranged from 62 to 89% (4.7 to 10 kgha^{minus 1} yr^{minus 1}) of annual inputs. DON made up themajority of TDN in stream exports, suggesting thatinclusion of DON is critical to assessing N dynamicseven in areas with large anthropogenic inputs of DIN.Despite the dominance of DON in streamwater,precipitation inputs of DON were approximately equalto outputs. DON concentrations in streamwater did notappear significantly influenced by seasonal biologicalcontrols, but did increase with discharge on somewatersheds. Streamwater NO₃-N was the onlyfraction of N that exhibited a seasonal pattern, withconcentrations increasing during the winter months andpeaking during snowmelt runoff. Concentrations ofNO₃-N varied considerably among watersheds andare related to DOC:DON ratios in streamwater. AnnualDIN exports were negatively correlated withstreamwater DOC:DON ratios, indicating that theseratios might be a useful index of N status of uplandforests.     

The effect of land use on dissolved organic carbon and nitrogen uptake in streams

1. Agricultural and urban land use may increase dissolved inorganic nitrogen (DIN) concentrations in streams and saturate biotic nutrient demand, but less is known about their impacts on the cycling of organic nutrients. To assess these impacts we compared the uptake of DIN (as ammonium, NH₄⁺), dissolved organic carbon (DOC, as acetate), and dissolved organic nitrogen (DON, as glycine) in 18 low‐gradient headwater streams in southwest Michigan draining forested, agricultural, or urban land‐use types. Over 3 years, we quantified uptake in two streams in each of the three land‐use types during three seasons (spring, summer and autumn). 2. We found significantly higher NH₄⁺ demand (expressed as uptake velocity, V_f) in urban compared to forested streams and NH₄⁺V_f was greater in spring compared to summer and autumn. Acetate V_f was significantly higher than NH₄⁺ and glycine V_f, but neither acetate nor glycine V_f were influenced by land‐use type or season. 3. We examined the interaction between NH₄⁺ and acetate demand by comparing simultaneous short‐term releases of both solutes to releases of each solute individually. Acetate V_f did not change during the simultaneous release with NH₄⁺, but NH₄⁺V_f was significantly higher with increased acetate. Thus, labile DOC V_f was not limited by the availability of NH₄⁺, but NH₄⁺V_f was limited by the availability of labile DOC. In contrast, neither glycine nor NH₄⁺V_f changed when released simultaneously indicating either that overall N‐uptake was saturated or that glycine and NH₄⁺ uptake were controlled by different factors. 4. Our results suggest that labile DOC and DON uptake can be equivalent to, or even higher than NH₄⁺ uptake, a solute known to be highly bioreactive, but unlike NH₄⁺ uptake, may not differ among land‐use types and seasons. Moreover, downstream export of nitrogen may be exacerbated by limitation of NH₄⁺ uptake by the availability of labile DOC in headwater streams from the agricultural Midwestern United States. Further research is needed to identify the factors that influence cycling of DOC and DON in streams.     

陆地生态系统可溶性有机氮研究进展

可溶性有机氮(dissolved organic nitrogen DON)的流动是陆地生态系统氮循环的重要组成部分。本文就陆地生态系统DON的来源、组成、性质;森林生态系统DON的流动、季节动态以及DON在氮循环中的地位等方面作了概括和探讨。今后的陆地生态系统DON的研究应该集中在以下几个方面：确定陆地生态系统中DON的各分室DON的浓度、流量;DON的源与汇问题;量化不同生态系统中DON库的大小和组成;研究DON在氮的矿化、微生物的固持、以及植物吸收等氮循环过程中的地位;对比研究DON与DOC(dissolved organic carbon)的动态差别;探讨DON与植物营养和碳积累的关系等。     

Dynamics of dissolved organic nitrogen in subalpine Castle Lake,California

Dissolved organic nitrogen (DON) concentrations were measured in meso-oligotrophic Castle Lake, California during the ice-free season in 1982, 1983 and 1984. No consistent relationships were found between DON and particulate-N, primary productivity rates or chlorophyll concentrations. However, increases in DON concentrations were observed in the early growing season of 1982 and 1984 when water temperature was rising and a diatom bloom was senescing. DON increased at a high rate (0.31 mg atom N m^–3 day^–1), and then rapidly disappeared. Sediment released appared to be the most important source of DON. Dissolved free amino acids were always less than 7.5% of the DON pool, and did not vary in the composition of specific amino acids during the growing season.