Ecosystem thresholds with hypoxia

Hypoxia is one of the common effects of eutrophication in coastal marine ecosystems and is becoming an increasingly prevalent problem worldwide. The causes of hypoxia are associated with excess nutrient inputs from both point and non-point sources, although the response of coastal marine ecosystems is strongly modulated by physical processes such as stratification and mixing. Changes in climate, particularly temperature, may also affect the susceptibility of coastal marine ecosystems to hypoxia. Hypoxia is a particularly severe disturbance because it causes death of biota and catastrophic changes in the ecosystem. Bottom water oxygen deficiency not only influences the habitat of living resources but also the biogeochemical processes that control nutrient concentrations in the water column. Increased phosphorus fluxes from sediments into overlying waters occur with hypoxia. In addition, reductions in the ability of ecosystems to remove nitrogen through denitrification and anaerobic ammonium oxidation may be related to hypoxia and could lead to acceleration in the rate of eutrophication. Three large coastal marine ecosystems (Chesapeake Bay, Northern Gulf of Mexico, and Danish Straits) all demonstrate thresholds whereby repeated hypoxic events have led to an increase in susceptibility of further hypoxia and accelerated eutrophication. Once hypoxia occurs, reoccurrence is likely and may be difficult to reverse. Therefore, elucidating ecosystem thresholds of hypoxia and linking them to nutrient inputs are necessary for the management of coastal marine ecosystems. Finally, projected increases in warming show an increase in the susceptibility of coastal marine ecosystems to hypoxia such that hypoxia will expand. Guest editors: J. H. Andersen & D. J. Conley Eutrophication in Coastal Ecosystems: Selected papers from the Second International Symposium on Research and Management of Eutrophication in Coastal Ecosystems, 20–23 June 2006, Nyborg, Denmark     

Nitrogen budgets for the Republic of Korea and the Yellow Sea region

Growing populations in northeast Asia have greatly altered the nitrogencycle, with increases in agricultural production to feed the population, andwith increases in N emissions and transboundary air pollution. For example,during the 1900's over 50% of the N deposition over Republic of Korea wasimported from abroad. In this paper, we present biogeochemical budgets ofN for the South Korean peninsula (the Republic of Korea) and for the YellowSea region. We quantify N inputs from atmospheric deposition, fertilizers,biological fixation, and imports of food, feed, and products. We quantifyoutputs in riverine export, crop uptake, denitrification, volatilization,runoff, sedimentation and sea water exchange. Calculations were conductedusing mean values from 1994–1997. All of the nitrogen budgets werepositive, with N inputs exceeding outputs. The excess N inputs gave rise toincreases in N storage in landfills and in groundwater. Annual accumulationof N in the Yellow sea, including inputs from South Korea and otherdrainage areas, was 1229 kt yr^–1 with a residence time for N ofapproximately 1.5 years, thus doubling N content in marine waters every 3years during 1994–1997. The human derived N inputs leads to excessiveeutrophication and pollution of the Yellow Sea.     

Denitrification in coastal ecosystems: methods, environmental controls, and ecosystem level controls, a review

In this review of sediment denitrification in estuaries and coastal ecosystems, we examine current denitrification measurement methodologies and the dominant biogeochemical controls on denitrification rates in coastal sediments. Integrated estimates of denitrification in coastal ecosystems are confounded by methodological difficulties, a lack of systematic understanding of the effects of changing environmental conditions, and inadequate attention to spatial and temporal variability to provide both seasonal and annual rates. Recent improvements in measurement techniques involving ¹⁵ N techniques and direct N₂ concentration changes appear to provide realistic rates of sediment denitrification. Controlling factors in coastal systems include concentrations of water column NO ₃ ^– , overall rates of sediment carbon metabolism, overlying water oxygen concentrations, the depth of oxygen penetration, and the presence/absence of aquatic vegetation and macrofauna. In systems experiencing environmental change, either degradation or improvement, the importance of denitrification can change. With the eutrophication of the Chesapeake Bay, the overall rates of denitrification relative to N loading terms have decreased, with factors such as loss of benthic habitat via anoxia and loss of submerged aquatic vegetation driving such effects.     

Biotic versus hydrologic control over seasonal nitrate leaching in a floodplain forest

Strong seasonal increases in aquatic (stream, ground and hyporheic water) nitrate have been observed in a variety of ecosystems. In most cases, changes in hydrological and vegetative activity occur contemporaneously, making it difficult to determine whether soil leaching is being driven by increases in the availability of leachable N or is simply due to flushing of N that has accumulated over longer periods. Three studies were conducted to better determine controls on soil nitrate leaching in a near-pristine temperate floodplain ecosystem receiving large N inputs via N-fixation by red alder: 1) an artificial rainfall experiment was conducted to estimate N-leaching potential during the summer, when plant uptake is high and new inputs of organic matter are low; 2) soil solution, groundwater and surface water were sampled during a major autumn storm to document exchanges at the seasonal transition, when plant uptake is low and inputs of senescent organic matter are high; and 3) monthly samples of soil and aquatic nitrogen were collected in 1997 and 1998 to document seasonal patterns of N exchanges. Collectively, these studies demonstrate the importance of hydrologic factors in controlling N flux. Nitrate was rapidly leached from soils during actual and simulated rainstorms. Two pathways of nitrate leaching were identified. Localized flooding and direct leaching of streamside soils into surface waters contributed to high solute concentrations in peak flows. Nitrate that leached into interstitial waters was subject to various factors that could delay or reduce its delivery to surface waters. Greater residence time may increase the influence of this component of stormflow on ecosystem productivity. While soil nitrate pools were rapidly depleted during rainstorms, accumulation of soil nitrate occurred over summer dry periods. Large differences in soil and aquatic nitrate concentrations between two years with contrasting rainfall highlight the potential for inter-annual hydrologic variability to affect ecosystem nutrient cycling.     

A review of some characteristics of freshwater and coastal ecosystems in Ghana

Data collected over a ten year period have been used to review some characteristics of freshwater and coastal ecosystems in Ghana. Studies were conducted on two recently formed man-made lakes, one river, five coastal lagoons and waters along the ocean front.Freshwaters had near neutral pH while coastal waters were alkaline. In both ecosystems the pH of sediments were lower than those of corresponding waters. While the transparency of coastal waters varied within narrow limits, large variations were observed in freshwaters. The lowest concentration of nutrients occurred in the river and the highest in coastal waters.In general, differences in these physical and chemical characteristics do not depend on whether the aquatic systems were fresh, brackish or saline. They seem to be more influenced by such factors as whether a water body: (1) was lotic or lentic, (2) was influenced by industrial or domestic activities, (3) was affected by impoundment, (4) received turbid water inflows, or (5) was eutrophic.     

Thaw depth determines reaction and transport of inorganic nitrogen in valley bottom permafrost soils

Nitrate (NO₃^–) export coupled with high inorganic nitrogen (N) concentrations in Alaskan streams suggests that N cycles of permafrost‐influenced ecosystems are more open than expected for N‐limited ecosystems. We tested the hypothesis that soil thaw depth governs inorganic N retention and removal in soils due to vertical patterns in the dominant N transformation pathways. Using an in situ, push–pull method, we estimated rates of inorganic N uptake and denitrification during snow melt, summer, and autumn, as depth of soil–stream flowpaths increased in the valley bottom of an arctic and a boreal catchment. Net NO₃^– uptake declined sharply from snow melt to summer and decreased as a nonlinear function of thaw depth. Peak denitrification rate occurred during snow melt at the arctic site, in summer at the boreal site, and declined as a nonlinear function of thaw depth across both sites. Seasonal patterns in ammonium (NH₄⁺) uptake were not significant, but low rates during the peak growing season suggest uptake that is balanced by mineralization. Despite rapid rates of hydrologic transport during snow melt runoff, rates of uptake and removal of inorganic N tended to exceed water residence time during snow melt, indicating potential for retention of N in valley bottom soils when flowpaths are shallow. Decreased reaction rates relative to water residence time in subsequent seasons suggest greater export of inorganic N as the soil–stream flowpath deepens due to thawing soils. Using seasonal thaw as a proxy for longer term deepening of the thaw layer caused by climate warming and permafrost degradation, these results suggest increasing potential for export of inorganic N from permafrost‐influenced soils to streams.     

Ecological risks in anthropogenic disturbance of nitrogen cycles in natural terrestrial ecosystems

Anthropogenic addition of reactive nitrogen (Nr) to the biosphere is increasing globally and some terrestrial ecosystems are suffering from a state of excess Nr for biological nitrogen (N) demand, termed N saturation. Here, we review the ecological risks in relation to N saturation and prospective responses to N saturation. Excess Nr increases the risks of local extinction of rare plant species, encouragement of exotic plant species, disturbance of nutrient balance in plant organs, and increase of herbivory in plant communities. On the ecosystem scale, excess bioavailable N induces forest decline, disturbance of nutrient cycling within ecosystems, depending on vegetation, soil, land-use, and N-loading history. These Nr risks will increase in the Asian region, where impacts of Nr in natural terrestrial ecosystems have been scarcely studied. Whether much of the terrestrial ecosystems on a global level are in the sate of N saturation or not is still controversial, but the potential risks of excess Nr seem to be increasing. The fundamental ways to mitigate Nr risks are to reduce Nr production, prevent Nr translocation, and promote conversion of Nr to N₂. Temporal, but promising actions against ecological N risks may include management of forests and riparian zones, and carbon addition in grassland.     

Nitrogen cycling and anthropogenic impact in the tropical interamerican seas

We discuss the mechanisms leading to nutrient limitation in tropical marine systems, with particular emphasis on nitrogen cycling in Caribbean ecosystems. We then explore how accelerated nutrient cycling from human activities is affecting these systems.Both nitrogen and phosphorus exert substantial influence on biological productivity and structure of tropical marine ecosystems. Offshore planktonic communities are largely nitrogen limited while nearshore ecosystems are largely phosphorus limited. For phosphorus, the ability of sediment to adsorb and store phosphorus is probably greater for tropical carbonate sediments than for most nearshore sediments in temperate coastal systems. However, the ability of tropical carbonate sediments to take up phosphorus can become saturated as phosphorus loading from human sources increases. The nature of the sediment, the mixing rate between nutrient-laden runoff waters and nutrient-poor oceanic waters and the degree of interaction of these water masses with the sediment will probably control the dynamics of this transition.Nearshore tropical marine ecosystems function differently from their temperate counterparts where coupled nitrification/denitrification serves as an important mechanism for nitrogen depuration. In contrast, nearshore tropical ecosystems are more susceptible to nitrogen loading as depurative capacity of the microbial communities is limited by the fragility of the nitrification link. At the same time, accumulation of organic matter in nearshore carbonate sediments appears to impair their capacity for phosphorus immobilization. In the absence of depurative mechanisms for either phosphorus or nitrogen, limitation for both these nutrients is alleviated and continued nutrient loading fuels the proliferation of nuisance algae.     

Global pollution of surface waters from point and nonpoint sources of nitrogen

Global 0.5- by 0.5-degree resolution estimates are presented on the fate of nitrogen (N) stemming from point and nonpoint sources, including plant uptake, denitrification, leaching from the rooting zone, rapid flow through shallow groundwater, and slow flow through deep groundwater to riverine systems. Historical N inputs are used to describe the N flows in groundwater. For nonpoint N sources (agricultural and natural ecosystems), calculations are based on local hydrology, climate, geology, soils, climate and land use combined with data for 1995 on crop production, N inputs from N fertilizers and animal manure, and estimates for ammonia emissions, biological N fixation, and N deposition. For point sources, our estimates are based on population densities and human N emissions, sanitation, and treatment. The results provide a first insight into the magnitude of the N losses from soil-plant systems and point sources in various parts of the world, and the fate of N during transport in atmosphere, groundwater, and surface water. The contribution to the river N load by anthropogenic N pollution is dominant in many river basins in Europe, Asia, and North Africa. Our model results explain much of the variation in measured N export from different world river basins.     

Transport of groundwater-borne nutrients from watersheds and their effects on coastal waters

Anthropogenic activities on coastal watersheds increase nutrient concentrations of groundwater. As groundwater travels downslope it transports these nutrients toward the adjoining coastal water. The resulting nutrient loading rates can be significant because nutrient concentrations in coastal groundwaters may be several orders of magnitude greater than those of receiving coastal waters. Groundwater-borne nutrients are most subject to active biogeochemical transformations as they course through the upper 1 m or so of bottom sediments. There conditions favor anaerobic processes such as denitrification, as well as other mechanisms that either sequester or release nutrients. The relative importance of advective vs. regenerative pathways of nutrient supply may result in widely different rates of release of nutrients from sediments. The relative activity of denitrifiers also may alter the ratio of N to P released to overlying waters, and hence affect which nutrient limits growth of producers. The consequences of nutrient (particularly nitrate) loading include somewhat elevated nutrient concentrations in the watercolumn, increased growth of macroalgae and phytoplankton, reduction of seagrass beds, and reductions of the associated fauna. The decline in animals occurs because of habitat changes and because of the increased frequency of anoxic events prompted by the characteristically high respiration rates found in enriched waters.     

Transport of groundwater-borne nutrients from watersheds and their effects on coastal waters

Anthropogenic activities on coastal watersheds increase nutrient concentrations of groundwater. As groundwater travels downslope it transports these nutrients toward the adjoining coastal water. The resulting nutrient loading rates can be significant because nutrient concentrations in coastal groundwaters may be several orders of magnitude greater than those of receiving coastal waters. Groundwater-borne nutrients are most subject to active biogeochemical transformations as they course through the upper 1 m or so of bottom sediments. There conditions favor anaerobic processes such as denitrification, as well as other mechanisms that either sequester or release nutrients. The relative importance of advective vs. regenerative pathways of nutrient supply may result in widely different rates of release of nutrients from sediments. The relative activity of denitrifiers also may alter the ratio of N to P released to overlying waters, and hence affect which nutrient limits growth of producers. The consequences of nutrient (particularly nitrate) loading include somewhat elevated nutrient concentrations in the watercolumn, increased growth of macroalgae and phytoplankton, reduction of seagrass beds, and reductions of the associated fauna. The decline in animals occurs because of habitat changes and because of the increased frequency of anoxic events prompted by the characteristically high respiration rates found in enriched waters.     

Spatial Heterogeneity of Denitrification in Semi-Arid Floodplains

Riparian ecosystems are recognized as sinks for inorganic nitrogen (N). Denitrification, a heterotrophic microbial process, often accounts for a significant fraction of the N removed. Characteristics of both riparian soils and hydrologic vectors may constrain the locations where denitrification can occur within riparian ecosystems by influencing the distribution of substrates, water, and suitable redox conditions. We employed spatially explicit methods to quantify heterogeneity of soil characteristics and potential rate of denitrification in semi-arid riparian ecosystems. These results allow us to evaluate the relative contributions of hydrologic vectors (water courses that convey materials) and soil resources (materials required by biota) to spatial heterogeneity of denitrification. During dry and monsoon seasons we contrasted a mesic site, characterized by shallow groundwater and annual inundation by floods, with a xeric site that is inundated less often and has a deeper water table. Potential denitrification was detected throughout the mesic floodplain and the average rate of denitrification was greater at the mesic site than at the xeric site, indicating the influence of water availability on denitrification. At the xeric reach, sharp declines in pools of soil resources and rate of denitrification occurred away from the stream, demonstrating the importance of the stream in determining spatial patterns. Using geographically weighted regression analysis, we determined that soil organic matter and soil nitrate were significant predictors of denitrification at the xeric site, but that factors influencing denitrification varied spatially. Spatial heterogeneity of carbon (C) and N substrates in soils likely influenced spatial patterns of denitrification, but distribution of C and N substrates was ultimately organized by hydrologic vectors. Droughts will increase the abundance of reaches with hydrogeomorphic templates similar to the xeric reach studied here. Consequences of such a transition may include a reduced rate of denitrification and patchy distribution of denitrification in floodplain soils, which will decrease the contribution of riparian ecosystems to N removal. TKH designed and completed the study and wrote the paper; EAW contributed methods and edited the paper; NBG designed the study and edited the paper.     

Analysis of nitrogen saturation potential in Rocky Mountain tundra and forest: implications for aquatic systems

We employed grass and forest versions of the CENTURY model under a range of N deposition values (0.02–1.60 g N m^–2 y^–1) to explore the possibility that high observed lake and stream N was due to terrestrial N saturation of alpine tundra and subalpine forest in Loch Vale Watershed, Rocky Mountain National Park, Colorado. Model results suggest that N is limiting to subalpine forest productivity, but that excess leachate from alpine tundra is sufficient to account for the current observed stream N. Tundra leachate, combined with N leached from exposed rock surfaces, produce high N loads in aquatic ecosystems above treeline in the Colorado Front Range. A combination of terrestrial leaching, large N inputs from snowmelt, high watershed gradients, rapid hydrologic flushing and lake turnover times, and possibly other nutrient limitations of aquatic organisms constrain high elevation lakes and streams from assimilating even small increases in atmospheric N. CENTURY model simulations further suggest that, while increased N deposition will worsen the situation, nitrogen saturation is an ongoing phenomenon.     

Interactive effects of anthropogenic nitrogen enrichment and climate change on terrestrial and aquatic biodiversity

Biodiversity has been described as the diversity of life on earth within species, among species, and among ecosystems. The rate of biodiversity loss due to human activity in the last 50 years has been more rapid than at any other time in human history, and many of the drivers of biodiversity loss are increasing, including habitat loss, overexploitation, invasive species, climate change, and pollution, including pollution from reactive nitrogen (Nr). Of these stressors, climate change and Nr from anthropogenic activities are causing some of the most rapid changes. Climate change is causing warming trends that result in poleward and elevational range shifts of flora and fauna, and changes in phenology, particularly the earlier onset of spring events and migration, and lengthening of the growing season. Nitrogen (N) enrichment can enhance plant growth, but has been shown to favor, fast-growing, sometimes invasive, species over native species adapted to low N conditions. Although there have been only a few controlled studies on climate change and N interactions, inferences can be drawn from various field observations. For example, in arid ecosystems of southern California, elevated N deposition and changing precipitation patterns have promoted the conversion of native shrub communities to communities dominated by annual non-native grasses. Both empirical studies and modeling indicate that N and climate change can interact to drive losses in biodiversity greater than those caused by either stressor alone. Reducing inputs of anthropogenic Nr may be an effective mitigation strategy for protecting biodiversity in the face of climate change.     

The Ratios of Carbon,Nitrogen, and Phosphorus in a Wetland Coastal Ecosystem of Southern India

The fertility of the coastal and estuarine waters is of great concern because of its influence on the productivity of these waters. Seasonal variations in the distribution of organic carbon, total nitrogen and total phosphorus in the sediments of Kuttanad Waters, a part of the tropical Cochin Estuary on the south west coast of India, are examined to identify the contribution of sediments to the fertility of the aquatic systems. The adjoining region has considerable agricultural activity. The fresh water zones had higher quantities of silt and clay whereas the estuarine zone was more sandy. Organic carbon, total phosphorus and total nitrogen were higher in the fresh water zones and lower in the estuarine zones. Total phosphorus and organic carbon showed the lowest values during monsoon periods. No significant trends were observed in the seasonal distributions of total nitrogen. Ratios of C/N, C/P and N/P, and the phosphorus and nitrogen content indicate significant modification in the character of the organic matter. Substantial amounts of the organic matter can contribute to reducing conditions and modify diagenetic processes.     

Research Commentary: Association of Zoonotic Pathogens with Fresh, Estuarine, and Marine Macroaggregates

Aquatic macroaggregates (flocs ≥0.5 mm) provide an important mechanism for vertical flux of nutrients and organic matter in aquatic ecosystems, yet their role in the transport and fate of zoonotic pathogens is largely unknown. Terrestrial pathogens that enter coastal waters through contaminated freshwater runoff may be especially prone to flocculation due to fluid dynamics and electrochemical changes that occur where fresh and marine waters mix. In this study, laboratory experiments were conducted to evaluate whether zoonotic pathogens (Cryptosporidium, Giardia, Salmonella) and a virus surrogate (PP7) are associated with aquatic macroaggregates and whether pathogen aggregation is enhanced in saline waters. Targeted microorganisms showed increased association with macroaggregates in estuarine and marine waters, as compared with an ultrapure water control and natural freshwater. Enrichment factor estimations demonstrated that pathogens are 2–4 orders of magnitude more concentrated in aggregates than in the estuarine and marine water surrounding the aggregates. Pathogen incorporation into aquatic macroaggregates may influence their transmission to susceptible hosts through settling and subsequent accumulation in zones where aggregation is greatest, as well as via enhanced uptake by invertebrates that serve as prey for marine animals or as seafood for humans.     

Comparison of denitrification characteristics among three habitat types of a large river floodplain: Atchafalaya River Basin,Louisiana

Mobile forms of nitrogen leach from upland environments into aquatic systems, often discharging to coastal zones. Addition of nitrogen to once N-limited systems results in a host of changes ranging from eutrophication to loss of biodiversity. Floodplains can ameliorate these changes by removing and sequestering nitrogen. In many coastal floodplains, sedimentation causes lakes to transition to baldcypress swamps, and ultimately to bottomland hardwood (BLHW) forests. These habitats differ in their contact with floodwater, which directly and indirectly affects their ability to process nutrients, but the effects of habitat change on denitrification at the floodplain scale cannot be predicted because of lack of suitable data. This study compared denitrification characteristics among the aforementioned habitats within the Atchafalaya River Basin (ARB). Microcosms were established in the laboratory, and the acetylene block technique was used to estimate four denitrification characteristics: background denitrification rates, maximum rates, time to reach maximum rates, and the linear response of denitrification to nitrate concentration. There were significant differences in denitrification characteristics among the three habitat types; specifically, all habitats differed in the time required for denitrification to respond to nitrate in the overlying water, and denitrification in lake habitats differed from both BLHWs and baldcypress swamps. Landscape-scale models should account for different linear relationships between denitrification and nitrate concentrations, and different response times to nitrate concentrations for different habitats. Because denitrification characteristics differ across habitats within the ARB, continued habitat change within the floodplain will alter nutrient discharge to coastal waters.     

The biogeochemistry of nitrogen in freshwater wetlands

The biogeochemistry of N in freshwater wetlands is complicated by vegetation characteristics that range from annual herbs to perennial woodlands; by hydrologic characteristics that range from closed, precipitation-driven to tidal, riverine wetlands; and by the diversity of the nitrogen cycle itself. It is clear that sediments are the single largest pool of nitrogen in wetland ecosystems (100's to 1000's g N m^-2) followed in rough order-of-magnitude decreases by plants and available inorganic nitrogen. Precipitation inputs (< 1–2 g N m^-2 yr^-1) are well known but other atmospheric inputs, e.g. dry deposition, are essentially unknown and could be as large or larger than wet deposition. Nitrogen fixation (acetylene reduction) is an important supplementary input in some wetlands (< < 1–3 g N m^-2 yr^-1) but is probably limited by the excess of fixed nitrogen usually present in wetland sediments.Plant uptake normally ranges from a few g N m^-2 yr^-1 to 35 g N m^-2 yr^-1 with extreme values of up to 100g N m^-2 yr^-1 Results of translocation experiments done to date may be misleading and may call for a reassessment of the magnitude of both plant uptake and leaching rates. Interactions between plant litter and decomposer microorganisms tend, over the short-term, to conserve nitrogen within the system in immobile forms. Later, decomposers release this nitrogen in forms and at rates that plants can efficiently reassimilate.The NO₃ formed by nitrification (< 0.1 to 10 g N m^-2 yr^-1 has several fates which may tend to either conserve nitrogen (uptake and dissimilatory reduction to ammonium) or lead to its loss (denitrification). Both nitrification and denitrification operate at rates far below their potential and under proper conditions (e.g. draining or fluctuating water levels) may accelerate. However, virtually all estimates of denitrification rates in freshwater wetlands are based on measurements of potential denitrification, not actual denitrification and, as a consequence, the importance of denitrification in these ecosystems may have been greatly over estimated.In general, larger amounts of nitrogen cycle within freshwater wetlands than flow in or out. Except for closed, ombrotrophic systems this might seem an unusual characteristic for ecosystems that are dominated by the flux of water, however, two factors limit the opportunity for N loss. At any given time the fraction of nitrogen in wetlands that could be lost by hydrologic export is probably a small fraction of the potentially mineralizable nitrogen and is certainly a negligible fraction of the total nitrogen in the system. Second, in some cases freshwater wetlands may be hydrologically isolated so that the bulk of upland water flow may pass under (in the case of floating mats) or by (in the case of riparian systems) the biotically active components of the wetland. This may explain the rather limited range of N loading rates real wetlands can accept in comparison to, for example, percolation columns or engineered marshes.     

A case study of nitrogen saturation in western U.S. forests

Virtually complete nitrification of the available ammonium in soil and nitrification activity in the forest floor are important factors predisposing forests in the San Bernardino Mountains of southern California to nitrogen (N) saturation. As a result, inorganic N in the soil solution is dominated by nitrate. High nitrification rates also generate elevated nitric oxide (NO) emissions from soil. High-base cation saturation of these soils means that soil calcium depletion or effects associated with soil acidification are not an immediate risk for forest health as has been postulated for mesic forests in the eastern U.S. Physiological disturbance (e.g., altered carbon [C] cycling, reduced fine root biomass, premature needle abscission) of ozone-sensitive ponderosa pine trees exposed to high N deposition and high ozone levels appear to be the greater threat to forest sustainability. However, N deposition appears to offset the aboveground growth depression effects of ozone exposure. High nitrification activity reported for many western ecosystems suggests that with chronic N inputs these systems are prone to N saturation and hydrologic and gaseous losses of N. High runoff during the winter wet season in California forests under a Mediterranean climate may further predispose these watersheds to high nitrate leachate losses. After 4 years of N fertilization at a severely N saturated site in the San Bernardino Mountains, bole growth unexpectedly increased. Reduced C allocation below- ground at this site, presumably in response to ozone or N or both pollutants, may enhance the bole growth response to added N.