Nitrogen deposition in California forests: a review

Atmospheric concentrations and deposition of the major nitrogenous (N) compounds and their biological effects in California forests are reviewed. Climatic characteristics of California are summarized in light of their effects on pollutant accumulation and transport. Over large areas of the state dry deposition is of greater magnitude than wet deposition due to the arid climate. However, fog deposition can also be significant in areas where seasonal fogs and N pollution sources coincide. The dominance of dry deposition is magnified in airsheds with frequent temperature inversions such as occur in the Los Angeles Air Basin. Most of the deposition in such areas occurs in summer as a result of surface deposition of nitric acid vapor (HNO3) as well as particulate nitrate (NO3-) and ammonium (NH4+). Internal uptake of gaseous N pollutants such as nitrogen dioxide (NO2), nitric oxide (NO), HNO3, peroxyacetyl nitrate (PAN), ammonia (NH3), and others provides additional N to forests. However, summer drought and subsequent lower stomatal conductance of plants tend to limit plant utilization of gaseous N. Nitrogen deposition is much greater than S deposition in California. In locations close to photochemical smog source areas, concentrations of oxidized forms of N (NO2, HNO3, PAN) dominate, while in areas near agricultural activities the importance of reduced N forms (NH3, NH4+) significantly increases. Little data from California forests are available for most of the gaseous N pollutants. Total inorganic N deposition in the most highly-exposed forests in the Los Angeles Air Basin may be as high as 25-45 kg ha(-1) year(-1). Nitrogen deposition in these highly-exposed areas has led to N saturation of chaparral and mixed conifer stands. In N saturated forests high concentrations of NO3- are found in streamwater, soil solution, and in foliage. Nitric oxide emissions from soil and foliar N:P ratios are also high in N saturated sites. Further research is needed to determine the ecological effects of chronic N deposition, and to develop appropriate management options for protecting water quality and managing plant nutrient resources in ecosystems which no longer retain excess N.     

Nutrient status and plant growth effects of forest soils in the Basin of Mexico

The nutrient status of forest soils in the Mexico City Air Basin was evaluated by observing plant growth responses to fertilization with N, P or both nutrients combined. P deficiency was the most frequent condition for soil from two high pollution sites and N deficiency was greatest at a low N deposition site. Concentrations of Pb and Ni, and to a lesser extent Zn and Co, were higher at the high pollution sites. However, positive plant growth responses to P and sometimes to N, and results of wheat root elongation bioassays, suggest that heavy metal concentrations were not directly phytotoxic. Further studies are needed to determine if heavy metal toxicity to mycorrhizal symbionts of eucalyptus (Eucalyptus camaldulensis Dehnh.) from high pollution sites may explain the P deficiency and stunted growth. P deficiency is expected to limit the capacity for biotic N retention in N saturated forested watersheds in the Basin of Mexico dominated by Andisols.     

Nitrogen deposition effects on Mediterranean-type ecosystems: an ecological assessment

We review the ecological consequences of N deposition on the five Mediterranean regions of the world. Seasonality of precipitation and fires regulate the N cycle in these water-limited ecosystems, where dry N deposition dominates. Nitrogen accumulation in soils and on plant surfaces results in peaks of availability with the first winter rains. Decoupling between N flushes and plant demand promotes losses via leaching and gas emissions. Differences in P availability may control the response to N inputs and susceptibility to exotic plant invasion. Invasive grasses accumulate as fuel during the dry season, altering fire regimes. California and the Mediterranean Basin are the most threatened by N deposition; however, there is limited evidence for N deposition impacts outside of California. Consequently, more research is needed to determine critical loads for each region and vegetation type based on the most sensitive elements, such as changes in lichen species composition and N cycling.     

Effects of Atmospheric Nitrogen Deposition on Remote Freshwater Ecosystems

We review known and hypothesized effects of nitrogen (N) deposition owing to human activities on the chemistry, organisms, and ecosystem processes of remote oligotrophic freshwaters. Acidification is the best-known effect of N deposition on water chemistry, but additional effects include increased nutrient availability and alteration of the balance between N and other nutrients. Our synthesis of the literature, framed in a comprehensive model for the effects of N deposition on natural ecosystems, shows that all these effects can reduce biological diversity and alter ecosystem processes in remote freshwaters. N deposition is projected to grow worldwide in the near future and will interact with other global changes. Present effects on these fragile ecosystems may be only early signs of more radical impacts ahead.     

Effects of taxonomy,sediment, and water column on C:N:P stoichiometry of submerged macrophytes in Yangtze floodplain shallow lakes,China

Carbon (C), nitrogen (N) and phosphorus (P) are the three most important essential elements limiting growth of primary producers. Submerged macrophytes generally absorb nutrients from sediments by root uptake. However, the C:N:P stoichiometric signatures of plant tissue are affected by many additional factors such as taxonomy, nutrient availability, and light availability. We first revealed the relative importance of taxonomy, sediment, and water column on plant C:N:P stoichiometry using variance partitioning based on partial redundancy analyses. Results showed that taxonomy was the most important factor in determining C:N:P stoichiometry, then the water column and finally the sediment. In this study, a significant positive relationship was found between community C concentration and macrophyte community biomass, indicating that the local low C availability in macrophytes probably was the main reason why submerged macrophytes declined in Yangtze floodplain shallow lakes. Based on our study, it is suggested that submerged macrophytes in Yangtze floodplain shallow lakes are primarily limited by low light levels rather than nutrient availability.

     

Interactive effects of nitrogen and phosphorus loadings on nutrient removal from simulated wastewater using Schoenoplectus validus in wetland microcosms

The concentrations of nutrients (N and P) in the wastewater and loading rate to the constructed wetlands may influence the nutrient removal from the secondary-treated municipal wastewater using wetland plants. Three loading rates of N (low 5.7, medium 34.3 and high 103 mg N d(-1)) and two of P (low 3.4 and high 17.1 mg P d(-1)) were studied in simulated secondary-treated municipal wastewater using Schoenoplectus validus (Vahl) A. L?ve & D. L?ve in the vertical free surface-flow wetland microcosms. After 70-d growth, there were significant interactive effects of N and P on the total, above-ground and root biomass. The below-ground biomass (rhizome and root) was negatively affected by the high N treatment. The tissue concentrations of N increased with an increase in N additions and decreased with an increase in P applications, whereas the tissue concentrations of P increased with an increase in P additions and decreased with an increase in N applications at the low P treatment, but increased at the high P treatment. Significant interactive effects of N and P loadings were found for the removal efficiencies of NH(4) and P, but not that of NO(x). The plant uptake, substrate storage and other losses (e.g. denitrification and formation of organic film) had similar contribution to N removal when N loading was relatively low. The P storage by substrate was the main contribution to P removal when P loading was high, but plant uptake was the major factor responsible for P removal when P loading was low and N loading was high. The high nutrient availability and optimum ratio of N:P are required to stimulate growth of S. validus, resulting in preferential allocation of resources to the above-ground tissues and enhancing the nutrient removal efficiencies, but the high N concentration in wastewater may hamper the growth of S. validus in constructed wetlands.     

Effects of atmospheric ammonia (NH3) on terrestrial vegetation: a review

At the global scale, among all N (nitrogen) species in the atmosphere and their deposition on to terrestrial vegetation and other receptors, NH3 (ammonia) is considered to be the foremost. The major sources for atmospheric NH3 are agricultural activities and animal feedlot operations, followed by biomass burning (including forest fires) and to a lesser extent fossil fuel combustion. Close to its sources, acute exposures to NH3 can result in visible foliar injury on vegetation. NH3 is deposited rapidly within the first 4-5 km from its source. However, NH3 is also converted in the atmosphere to fine particle NH4+ (ammonium) aerosols that are a regional scale problem. Much of our current knowledge of the effects of NH3 on higher plants is predominantly derived from studies conducted in Europe. Adverse effects on vegetation occur when the rate of foliar uptake of NH3 is greater than the rate and capacity for in vivo detoxification by the plants. Most to least sensitive plant species to NH3 are native vegetation > forests > agricultural crops. There are also a number of studies on N deposition and lichens, mosses and green algae. Direct cause and effect relationships in most of those cases (exceptions being those locations very close to point sources) are confounded by other environmental factors, particularly changes in the ambient SO2 (sulfur dioxide) concentrations. In addition to direct foliar injury, adverse effects of NH3 on higher plants include alterations in: growth and productivity, tissue content of nutrients and toxic elements, drought and frost tolerance, responses to insect pests and disease causing microorganisms (pathogens), development of beneficial root symbiotic or mycorrhizal associations and inter species competition or biodiversity. In all these cases, the joint effects of NH3 with other air pollutants such as all-pervasive O3 or increasing CO2 concentrations are poorly understood. While NH3 uptake in higher plants occurs through the shoots, NH4+ uptake occurs through the shoots, roots and through both pathways. However, NH4+ is immobile in the soil and is converted to NO3- (nitrate). In agricultural systems, additions of NO3- to the soil (initially as NH3 or NH4+) and the consequent increases in the emissions of N2O (nitrous oxide, a greenhouse gas) and leaching of NO3- into the ground and surface waters are of major environmental concern. At the ecosystem level NH3 deposition cannot be viewed alone, but in the context of total N deposition. There are a number of forest ecosystems in North America that have been subjected to N saturation and the consequent negative effects. There are also heathlands and other plant communities in Europe that have been subjected to N-induced alterations. Regulatory mitigative approaches to these problems include the use of N saturation data or the concept of critical loads. Current information suggests that a critical load of 5-10 kg ha(-1) year(-1) of total N deposition (both dry and wet deposition combined of all atmospheric N species) would protect the most vulnerable terrestrial ecosystems (heaths, bogs, cryptogams) and values of 10-20 kg ha(-1) year(-1) would protect forests, depending on soil conditions. However, to derive the best analysis, the critical load concept should be coupled to the results and consequences of N saturation.     

Plant senescence: a mechanism for nutrient release in temperate agricultural wetlands

The beneficial uptake of nutrients by wetland plants is countered to some extent by nutrient release back into the aquatic environment due to vegetative die-back. This current study examined whether Leersia oryzoides, a common wetland plant, exhibits luxury uptake of nutrients from simulated farm runoff. The study also tested whether with subsequent decomposition, these nutrients are released back into the water column. When exposed to elevated (>2mg/L N and P) runoff, L. oryzoides assimilated significantly higher concentrations of nitrogen (p<0.001) and phosphorus (p<0.001) in above-ground biomass as compared to non-enriched treatments (<0.05 mg/L N and P). Subsequently, senescence of enriched above-ground biomass yielded significantly higher concentrations of phosphorus (2.19+/-0.84 mg P/L). Using L. oryzoides as our model, this study demonstrates nitrogen and phosphorus sequestration during the growing season and release of phosphorus in the winter.     

Elemental composition of Juncus bulbosus in an acidified freshwater reservoir

Concentrations of the major nutrients (N, P, Mg, K, Na and trace elements (Fe, Mn, Zn) were determined in tissues of the submerged aquatic plant Juncus bulbosus L. collected from the acidified Freshwater Reservoir of Uusikaupunki in SW Finland. The heavy metals showed markedly elevated concentrations, mainly due to the enhanced solubility and availability of metals in the reservoir receiving acidic, sulphate-rich discharges from the catchment area. Tissue analyses of J. bulbosus could be used to reveal specific transport patterns in the reservoir. Multivariate principal components ordination of the data of the elemental analyses demonstrated local differences in the availability of metals.     

Effects of photochemical smog and mineral nutrition on ponderosa pine seedlings

Two-year-old seedlings of ponderosa pine (Pinus ponderosa Dougl. ex Laws) were exposed to ambient concentrations of photochemical smog (AA) and clean air (CA) during a single field season at Tanbark Flat of the San Gabriel Mountains in the Los Angeles Basin. The seedlings were grown in a perlite-vermiculite medium with full supply of nutrients (based on modified Hoagland solution); reduced to 50% supply of N; reduced to 50% supply of Mg; and reduced to 50% supply of N+Mg. No significant effects of air pollution exposures on injury development, stem growth and concentrations of plant pigments were determined. The seedlings in the AA treatment had decreased N concentration in current year needles compared with CA seedlings; however, the needle concentrations of other elements did not change. Reduction of N supply in the growing medium caused decreased N, P, Ca, K and chlorophyll a concentrations in needles. Stem growth of the seedlings with reduced N supply was significantly decreased as well. No changes in stem growth or chemical composition of plants with reduced Mg supply were noted. Reduction of supply of nutrients did not change responses of trees to the air pollution exposures.     

Use of Calluna vulgaris to detect signals of nitrogen deposition across an urban–rural gradient

Densely populated cities can experience high concentrations of traffic-derived pollutants, with oxides of nitrogen and ammonia contributing significantly to the overall nitrogen (N) budget of urban ecosystems. This study investigated changes in the biochemistry of in situ Calluna vulgaris plants to detect signals of N deposition across an urban–rural gradient from central London to rural Surrey, UK. Foliar N concentrations and δ¹⁵N signatures were higher, and C/N ratios lower, in urban areas receiving the highest rates of N deposition. Plant phosphorus (P) concentrations were also highest in these areas, suggesting that elevated rates of N deposition are unlikely to result in progressive P-limitation in urban habitats. Free amino acid concentrations were positively related to N deposition for asparagine, glutamine, glycine, phenylalanine, isoleucine, leucine and lysine. Overall, relationships between tissue chemistry and N deposition were similar for oxidised, reduced and total N, although the strength of relationships varied with the different biochemical indicators. The results of this study indicate that current rates of N deposition are having substantial effects on plant biochemistry in urban areas, with likely implications for the biodiversity and functioning of urban ecosystems.     

Functional Relationships with N Deposition Differ According to Stand Maturity in Calluna-Dominated Heathland

Plant and soil bio(chemical) indicators are increasingly used to provide information on N deposition inputs and effects in a wide range of ecosystem types. However, many factors, including climate and site management history, have the potential to influence bioindicator relationships with N due to nutrient export and changing vegetation nutrient demands. We surveyed 33 heathlands in England, along a gradient of background N deposition (7.2–24.5 kg ha⁻¹ year⁻¹), using Calluna vulgaris growth phase as a proxy for time since last management. Our survey confirmed soil nutrient accumulation with increasing time since management. Foliar N and phosphorus (P) concentrations in pioneer- and mature-phase vegetation significantly increased with N deposition. Significant interactions between climate and N deposition were also evident with, for example, higher foliar P concentrations in pioneer-phase vegetation at sites with higher temperatures and N deposition rates. Although oxidized N appeared more significant than reduced N, overall there were more, stronger relationships with total N deposition; suggesting efforts to control all emissions of N (i.e., both oxidized and reduced forms) will have ecological benefits.

Electronic supplementary material

The online version of this article (doi:10.1007/s13280-014-0529-4) contains supplementary material, which is available to authorized users.     

Effects of air filtration on spring wheat grown in open-top field chambers at a rural site. II. Effects on mineral partitioning, sulphur and nitrogen metabolism and on grain quality

In 1988 the effect of ambient levels of air pollutants on the nutrients status and grain quality of spring wheat (Triticum aestivum cv. Pelican) was investigated by comparing plants grown in open-top chambers (OTC) ventilated with ambient air (NF treatments) and charcoal-filtered air (CF treatments) at a rural site (Tervuren, Belgium). Spring wheat cultivated in NF OTC showed only minor differences in the P, K, Ca, Mg, Mn and Na concentrations of the different plant parts at final harvest, as well as organic and inorganic S fractions, compared to those of the plants grown in CF air. The plants' total P content was reduced, as well as the P and K concentration of the flour. The total S concentration of the flour was increased by 4%. Effects on N concentrations and grain quality were much more pronounced. At final harvest the N concentrations of straw and flour of the NF air treated plants were much higher compared to CF air. However, the N content of the aerial biomass and the grain N yield were not significantly affected, implying a reduction of other structual compounds. Nitrogen harvest index (NHI) and the ratio of NHI over grain harvest index (GHI), indicated a significant reduction of N translocation from the above-ground biomass to the grain. Changes in the N status and partitioning of spring wheat had an effect on the baking quality of wheat flour. Several parameters that are commonly used as an indication of baking quality have been significantly increased in the NF treatment: total protein concentration, Zeleny value, dry and wet gluten concentration. A slightly increased Hagberg value indicated a reduced alpha-amylase activity. The possibility of foliar N uptake as an additional N source, especially after anthesis and implications of increased protein production instead of carbohydrate synthesis are discussed.     

Can the foliar nitrogen concentration of upland vegetation be used for predicting atmospheric nitrogen deposition? Evidence from field surveys

The deposition of atmospheric nitrogen can be enhanced at high altitude sites as a consequence of cloud droplet deposition and orographic enhancement of wet deposition on hills. The degree to which the increased deposition of nitrogen influences foliar nitrogen concentration in a range of upland plant species was studied in a series of field surveys in northern Britain. A range of upland plant species sampled along altitudinal transects at sites of known atmospheric nitrogen deposition showed marked increases in foliar nitrogen concentration with increasing nitrogen deposition and altitude (and hence with decreasing temperature). For Nardus stricta L., Deschampsia flexuosa (L.) Trin., Calluna vulgaris (L.) Hull, Erica cinerea L. and Hylocomium splendens (Hedw.) Br. Eur. on an unpolluted hill, foliar nitrogen increased by 0.07, 0.12, 0.15, 0.08 and 0.04% dry weight respectively for each 1 kg ha(-1) year(-1) increase in nitrogen deposition. Most species showed an approximately linear relationship between foliar nitrogen concentration and altitude but no trend with altitude for foliar phosphorus concentration. This provided evidence that the tissue nutrient status of upland plants reflects nutrient availability rather than the direct effects of climate on growth. However, differences in the relationship between foliar nitrogen concentration and atmospheric nitrogen deposition for N. stricta sampled on hills in contrasting pollution climates show that the possibility of temperature-mediated growth effects on foliar nitrogen concentration should not be ignored. Thus, there is potential to use upland plant species as biomonitors of nitrogen deposition, but the response of different species to nitrogen addition, in combination with climatic effects on growth, must be well characterised.     

Metabolism of chlorpyrifos in relation to its effect on the availability of some plant nutrients in soil

A laboratory experiment was conducted to study the persistence and metabolism of chlorpyrifos in Gangetic Alluvial soil of West Bengal and also to evaluate their effect on the availability of the major plant nutrients (N, P and K) in soil following the application of chlorpyrifos @ 1 kg (T1), 10 kg (T2) and 100 kg (T3) a.i.ha(-1). The dissipation followed first order kinetics and the calculated half-life (T1/2) values ranged from 20 to 37 days. The primary metabolite of chlorpyrifos, 3,5,6-trichloropyridinol (TCP) was detected from 3rd day after application and was at maximum on 30th day which decreased progressively to non-detectable level (NDL) on 120th day for all the treatment doses. The secondary metabolite 3,5,6-trichloro-2-methoxy pyridine (TMP) was detected on 30th, 15th and 7th day in T1, T2 and T3 doses respectively which decreased to NDL during 90-120th day. ANOVA study revealed significant decrease in the available N and P content in soil treated with chlorpyrifos in comparison to the control set. The inhibitory effect on available N was attributable to TMP and for P it was due to the presence of TCP and TMP rather than chlorpyrifos itself as revealed by the step wise multiple regression technique. In the later stage of incubation, however the average N and P status was recovered significantly at 120 days which might be due to the disappearance of the metabolites. The variation due to time of observations or treatment doses was minimum in case of available K in soil.     

An integrative analysis of the role of atmospheric deposition and land management practices on nitrogen in the US agricultural sector

Additions of anthropogenic nitrogen (N) compounds constitute one of the major classes of air pollutants of significance to human health and the environment. Reliance on wet deposition measurements alone can lead to considerable underestimates (by 40-60%) of the total (wet + dry) atmospheric N deposition. In addition, wet deposition of N are about 20% of the levels that are lost due to volatilization (primarily ammonia). Nevertheless, in the agricultural sectors of the Mississippi River basins, farm management practices, and recycling of N within cropping systems clearly outweigh the contributions of atmospheric deposition. As opposed to native vegetation and forests, there are no records of the negative effects of atmospheric N deposition on crop yield. Similarly, field studies on the interactions of atmospheric N compounds with the incidence and spread of pathogens does not permit any generalizations. Nitrogen applied as fertilizer affects disease probably more by its effect on the plant growth than by its effects on pathogens. In contrast, atmospheric nitrogen dioxide appears to be a stimulant of aphid performance. Under conditions of heavy weed infestation, N fertilization stimulates weed growth and competitiveness, rather than crop yield.     

Bioavailability and plant accumulation of heavy metals and phosphorus in agricultural soils amended by long-term application of sewage sludge

Amendment of agricultural soils with municipal sewage sludges provides a valuable source of plant nutrients and organic matter. Nevertheless, addition of heavy metals and risks of eutrophication continue to be of concern. Metal behaviour in soils and plant uptake are dependent on the nature of the metal, sludge/soil physico-chemical properties and plant species. A pot experiment was carried out to evaluate plant production and heavy metal uptake, soil heavy metal pools and bioavailability, and soil P pools and possible leaching losses, in agricultural soils amended with sewage sludge for at least 10 years (F20) compared to non-amended soils (control). Sewage sludge application increased soil pH, N, Olsen-extractable-P, DOC and exchangeable Ca, Mg and K concentrations. Total and EDTA-extractable soil concentrations of Cu and Zn were also significantly greater in F20, and soil metal (Cu, Mn and Zn) and P fractionation altered. Compared to the control, in F20 relative amounts of acid-extractable (Mn, Zn), reducible (Mn, Zn) and oxidisable (Cu, Zn) metal fractions were greater, and a dominance of inorganic P forms was observed. Analyses of F20 soil solutions highlighted risks of PO4 and Cu leaching. However, despite the observed increases in metal bioavailability sewage sludge applications did not lead to an increase in plant shoot concentrations (in wild plants or crop species). On the contrary, depending on the plant species, Mn and Zn tissue concentrations were within the deficiency level for most plants.     

A laboratory investigation of the routes of HNO3 dry deposition to coniferous seedlings

White pine, Norway spruce and red spruce seedlings were exposed to nitric acid vapor concentrations of 10 to 120 ppb in constant stirred tank reactors. Nitric acid dry deposition rates were determined from both the change in nitric acid concentration in the reactor flow stream and from the amount of nitrogen recovered from the seedlings. Nitric acid labeled with 15N was used to distinguish dry-deposited nitrogen in the plant from the nitrogen that was already present. It was found that dry deposition occurs via three routes: surface deposition, trans-cuticular deposition, and stomatal deposition. Resistance to surface deposition is very low (< 4.8 m2-s mol(-1)) for a freshly washed surface, but increases as the surface adsorption sites are occupied. Resistance to trans-cuticular uptake averaged 206 m2-s mol(-1). Stomatal resistance can be calculated from the rate of water diffusion out of the plant. Eighty per cent of the nitric acid deposited via the trans-cuticular and stomatal routes was assimilated by the plant. However, none of the nitric acid deposited on the surface was assimilated. In rural areas with coniferous forests, the combination of low ambient nitric acid concentrations and low initial surface resistance means that most nitric acid will be dry deposited on the tree surface, and thus will not be directly assimilated.     

The impact of nitrogen deposition on acid grasslands in the Atlantic region of Europe

A survey of 153 acid grasslands from the Atlantic biogeographic region of Europe indicates that chronic nitrogen deposition is changing plant species composition and soil and plant-tissue chemistry. Across the deposition gradient (2-44 kg N ha⁻¹ yr⁻¹) grass richness as a proportion of total species richness increased whereas forb richness decreased. Soil C:N ratio increased, but soil extractable nitrate and ammonium concentrations did not show any relationship with nitrogen deposition. The above-ground tissue nitrogen contents of three plant species were examined: Agrostis capillaris (grass), Galium saxatile (forb) and Rhytidiadelphus squarrosus (bryophyte). The tissue nitrogen content of neither vascular plant species showed any relationship with nitrogen deposition, but there was a weak positive relationship between R. squarrosus nitrogen content and nitrogen deposition. None of the species showed strong relationships between above-ground tissue N:P or C:N and nitrogen deposition, indicating that they are not good indicators of deposition rate.