Nitrate reductase activity as an indicator of ponderosa pine response to atmospheric nitrogen deposition in the San Bernardino Mountains

Determinations of nitrate reductase (NR) activity in ponderosa pine (Pinus ponderosa Dougl. ex. Laws.) needles were performed during summer 1994 in two areas (consisting of six different sites) with different nitrogen (N) deposition levels in the San Bernardino Mountains, southern California. Nitrate reductase activity was used as an integrative indicator of atmospheric nitrogen deposition to pine trees (direct uptake of N species from the atmosphere and N transported from the soil). Deposition of nitrate (NO3-) to pine branches was measured in order to determine dry atmospheric inputs of the oxidized N species to tree foliage. High NR activity was detected in all of the experimental sites. Activity of the enzyme was significantly higher at the locations characterized by higher NO3- deposition to branches--slight positive correlation between branch deposited NO3- and NR activity was found. However, high variability of NR in time and between the experimental sites discredit the NR assay as a reliable indicator of N deposition for ponderosa pine in the field conditions. This could be caused by substantial interference from other abiotic and biotic factors with tropospheric ozone as probably the most important one.     

Relationships of ozone exposure to pine injury in the Sierra Nevada and San Bernardino Mountains of California, USA

Hourly ambient ozone exposure data and crown injury measurements were gathered in the Sierra Nevada and San Bernardino Mountains of California to develop relationships between the Ozone Injury Index (OII), the Forest Pest Management Index (FPM), chlorotic mottle, fascicle retention (OII index components) and cumulative ambient ozone indices for Pinus ponderosa Dougl. ex Laws and Pinus jeffreyi Grev. and Balf. Eleven sites located in the mixed conifer forest near ambient ozone monitoring sites were evaluated annually for 4 years. Four other sites in the San Bernardino Mountains were evaluated for 1 year. Analyses showed OII to be functionally equivalent (r2 = 0.96) to the FPM, and to depend only on fascicle retention and chlorotic mottle (R2 = 0.95) of the fourth whorl (or if four whorls are not present at the site, then the last whorl present for the majority of trees). Significant associations were found between OII and 4-year 24-h. summer SUM0, SUM06, W126 and HRS80 ozone indices. Three sites had higher levels of cumulative chlorotic mottle for individual whorls and larger numbers of trees with visible crown injury than other sites with similar cumulative ambient ozone levels. Including an indicator variable to discriminate between these two groups of sites increased R2 and decreased root mean square (RMSE) for all indices, especially SUM0 (R2 = 0.93, RMSE reduced by 46%).     

Dry deposition of sulfur dioxide or nitric acid to oak, elm and pine leaves

To improve our knowledge of atmospheric inputs to forests, experiments were performed in the field to measure the dry deposition of SO2 and HNO3 to oak, elm, and pine leaves. A tree branch was enclosed in a Teflon chamber, through which SO2 or HNO3 flowed. The dry deposition characteristics of SO2 and HNO3 were very different. The SO2 deposition occurred primarily through stomatal openings for the oak and pine leaves, and equal stomatal and cuticular deposition was observed for the elm leaves. The deposited SO2 could not be removed from the branch by extracting in water or by revolatilization. In contrast, over 90% of HNO3 dry deposition occurred to the cuticle. Most of the deposited HNO3 could be extracted from the leaves. Revolatilization of HNO3 was negligible from an active branch, but increased from a dormant or detached branch. A deposition velocity was derived from the ratio of the flux of the gas to the leaves and the gas concentration in the chamber. Deposition velocities ranged from 0.02 to 0.11 cm s(-1) for SO2 and from 0.2 to 1.2 cm s(-1) for HNO3 to individual leaf surfaces.     

Ozone distribution and phytotoxic potential in mixed conifer forests of the San Bernardino Mountains, southern California

In the San Bernardino Mountains of southern California, ozone (O(3)) concentrations have been elevated since the 1950s with peaks reaching 600ppb and summer seasonal averages >100ppb in the 1970s. During that period increased mortality of ponderosa and Jeffrey pines occurred. Between the late 1970s and late1990s, O(3) concentrations decreased with peaks approximately 180ppb and approximately 60ppb seasonal averages. However, since the late 1990s concentrations have not changed. Monitoring during summers of 2002-2006 showed that O(3) concentrations (2-week averages) for individual years were much higher in western sites (58-69ppb) than eastern sites (44-50ppb). Potential O(3) phytotoxicity measured as various exposure indices was very high, reaching SUM00 - 173.5ppmh, SUM60 - 112.7ppmh, W126 - 98.3ppmh, and AOT40 - 75ppmh, representing the highest values reported for mountain areas in North America and Europe.     

Contribution of canopy leaching to sulphate deposition in a Scots pine forest

Radioactive sulphate (35SO4) was applied to the soil below a Scots pine forest on 23 June 1989, and its movement into the canopy and into throughfall and stemflow was measured over 4 months. The specific activity, Bq (mg S)(-1), of the canopy increased monotonically; uptake by current-year (1989) expanding needles was initially twice as fast as by older needles or live twigs. By 10 October the canopy average specific activity was 62 Bq (mg S)(-1). The specific activity of net throughfall (throughfall + stemflow - rain), deduced from measurements from six throughfall collectors, six stemflow collectors and two rain collectors, fell rapidly from 12.6 Bq (mg S)(-1) in late July to <1 Bq (mg S)(-1) in mid-August. The results suggest (assuming rapid equilibration of 35S with sulphate in soil) that root-derived sulphate contributed c. 3% of sulphate in net throughfall and that dry deposition of SO2 and sulphate particles contributed c. 97% of the 0.56 g S m(-2) measured in net throughfall over the period. Simultaneous measurements of SO2 at canopy height and of NH3 above and within the canopy gave mean concentrations of 5.9 and 0.86 microg m(-3), respectively, sufficient to account for the sulphate measured in net throughfall only if codeposition of NH3 and SO2 occurred to canopy surfaces. The large values of specific activity observed in July, however, indicate that throughfall composition may be closely related to recent soil input of sulphate, and that equilibrium cannot be safely assumed. The possibility of a significant contribution of soil-derived sulphate to sulphate deposition in net throughfall cannot be ruled out on the basis of this experiment.     

Empirical and simulated critical loads for nitrogen deposition in California mixed conifer forests

Empirical critical loads (CL) for N deposition were determined from changes in epiphytic lichen communities, elevated NO(3)(-) leaching in streamwater, and reduced fine root biomass in ponderosa pine (Pinus ponderosa Dougl. ex Laws.) at sites with varying N deposition. The CL for lichen community impacts of 3.1kg ha(-1) year(-1) is expected to protect all components of the forest ecosystem from the adverse effects of N deposition. Much of the western Sierra Nevada is above the lichen-based CL, showing significant changes in lichen indicator groups. The empirical N deposition threshold and that simulated by the DayCent model for enhanced NO(3)(-)leaching were 17kg N ha(-1) year(-1). DayCent estimated that elevated NO(3)(-) leaching in the San Bernardino Mountains began in the late 1950s. Critical values for litter C:N (34.1), ponderosa pine foliar N (1.1%), and N concentrations (1.0%) in the lichen Letharia vulpina ((L.) Hue) are indicative of CL exceedance.     

Dry deposition of particles and canopy exchange: Comparison of wet,bulk and throughfall deposition at five forest sites in Italy

The contribution of dry deposition to the total atmospheric input of acidifying compounds and base cations is of overwhelming importance. Throughfall measurements provide an estimate of the total deposition to forest soils, including dry deposition, but some uncertainties, related to the canopy interaction processes, affect this approach. We compared the concentrations and the fluxes of the main ions determined in wet-only, bulk and throughfall samples collected at five forest sites in Italy. The contribution of coarse particles deposited onto the bulk samplers was of prime importance for base cations, representing on average from 16% to 46% of the bulk deposition. The extent of this dry deposition depended on some geographical features of the sites, such as the distance from the sea and the annual rainfall. The possibility of applying specific bulk/wet ratios to estimate the wet deposition proved to be limited by the temporal variability of these ratios, which must be considered together with the spatial variability. A direct comparison of the dry contribution deriving from the bulk–wet and the throughfall–wet demonstrated that an extensive natural surface (forest canopy) performs better than a small synthetic surface (funnel of the bulk sampler) in collecting dry deposition of SO₄²⁻, NO₃⁻ and Na⁺. The canopy exchange model was applied to both bulk and wet data to estimate the contribution of dry deposition to the total input of base cations, and the uncertainty associated to the model discussed. The exclusive use of bulk data led to a considerable underestimation of base cation dry deposition, which varies among the study sites.     

Measurement of dry deposition to surfaces in deciduous and pine canopies

The importance of dry deposition was assessed at perimeter and interior locations in two vegetative canopies. Dry deposition was measured directly by washing particles from leaves. Ambient particles and gases were also collected at both locations within the canopies. Ambient concentrations on the canopy interior were decreased relative to perimeter concentrations due to dry deposition scavenging by the canopy. The least scavenging was found for SO(4)(2-) and NH(4)(+) and the highest scavenging was found for HNO(3). Dry deposition of all species was higher to perimeter vegetative and surrogate surfaces than to interior surfaces, due both to the lower concentrations and the lower wind speeds in the sheltered interior. Deposition velocities compared well with other experimental and theoretical values.     

Dry deposition of airborne sulfate and nitrate to soybeans

Dry deposition contributes a substantial part of the total deposition of acidic pollutants and acid precursors to agricultural systems. However, because of the relative intractability of measurement of dry deposition fluxes, little work has been done to directly quantify dry inputs of pollutants to crops. In this research, foliar surface sampling ('leaf-washing') methods were developed and shown to be a practical and fairly precise means of monitoring the accumulation of dry-deposited SO4(2-) and NO3- on plant surfaces. Leaching of these ions from plant tissues was shown to be negligible; however, uptake by plants (e.g. stomatal gas exchange of SO2 or HNO3 and/or assimilation of surface accumulations of materials) is not accounted for by the sampling method. The significance of dry deposition to modification of the chemical microenvironment of leaf surfaces appears to be a factor of 3 to 20 or more greater than that of wet deposition alone. This is due to the cyclic reactivation of accumulated materials by dew and light rains, which may dissolve and mobilize, but not remove, the pollutant surface deposit. Therefore, while dry deposition of SO2 and SO4(2-) containing particles may contribute only part of the total mass of sulfur inputs to crop systems, the exposure of plant surface tissue to pollutants can be dominated by the dry-deposited material. The alteration of leaf surface chemistry may contribute to possible stress-producing mechanisms such as reduction of cuticular integrity, cellular injury and death, enhanced leaching of primary and secondary metabolites, and changes in pathogen infection efficiency.     

The effect of the limited availability of H2O2 in the competitive deposition of sulfur and oxidized nitrogen

To understand the influence of the reduction of SO2 emissions from a single source in the S and N deposition around its local environment, the application of the Sulphur Transport Eulerian Model 2 (STEM-II) was introduced in this paper. Observed local deposition patterns were analysed and explained in terms of the main processes involved in the pollutants deposition. It was necessary to take into account the limited availability of H2O2 because of its influence on both S(IV) and oxidized nitrogen deposition. In order to estimate the quantitative relationship between the SO2 emissions reduction and the observed S and N deposition patterns, these processes were simulated for different meteorological conditions. Simulation results were in agreement with both observed deposition patterns and limited availability of H2O2, specially if significant changes in the S deposition patterns were considered. Both observed and estimated S deposition patterns changed their top value location from the Southwest (1990) to the Northwest (1997) of the domain, because of the reduction of dry deposition. The global reduction of total S depositions, estimated and observed, were in good agreement too. Model simulations could explain the higher S dry deposition reduction, considering the emissions reduction strategy applied.     

Combined effects of nitrogen addition and organic matter manipulation on soil respiration in a Chinese pine forest

The response of soil respiration (Rs) to nitrogen (N) addition is one of the uncertainties in modelling ecosystem carbon (C). We reported on a long-term nitrogen (N) addition experiment using urea (CO(NH₂)₂) fertilizer in which Rs was continuously measured after N addition during the growing season in a Chinese pine forest. Four levels of N addition, i.e. no added N (N0: 0 g N m⁻² year⁻¹), low-N (N1: 5 g N m⁻² year⁻¹), medium-N (N2: 10 g N m⁻² year⁻¹), and high-N (N3: 15 g N m⁻² year⁻¹), and three organic matter treatments, i.e. both aboveground litter and belowground root removal (LRE), only aboveground litter removal (LE), and intact soil (CK), were examined. The Rs was measured continuously for 3 days following each N addition application and was measured approximately 3–5 times during the rest of each month from July to October 2012. N addition inhibited microbial heterotrophic respiration by suppressing soil microbial biomass, but stimulated root respiration and CO₂ release from litter decomposition by increasing either root biomass or microbial biomass. When litter and/or root were removed, the “priming” effect of N addition on the Rs disappeared more quickly than intact soil. This is likely to provide a point of view for why Rs varies so much in response to exogenous N and also has implications for future determination of sampling interval of Rs measurement.

     

Isozyme markers associated with O(3) tolerance indicate shift in genetic structure of ponderosa and Jeffrey pine in Sequoia National Park, California

Effects of canopy ozone (O(3)) exposure and signatures of genetic structure using isozyme markers associated with O(3) tolerance were analyzed in approximately 20-, approximately 80-, and >200-yr-old ponderosa (Pinus ponderosa Dougl. ex Laws.) and Jeffrey pine (Pinus jeffreyi Grev. & Balf.) in Sequoia National Park, California. For both species, the number of alleles and genotypes per loci was higher in parental trees relative to saplings. In ponderosa pine, the heterozygosity value increased, and the fixation index indicated reduction of homozygosity with increasing tree age class. The opposite tendencies were observed for Jeffrey pine. Utilizing canopy attributes known to be responsive to O(3) exposure, ponderosa pine was more symptomatic than Jeffrey pine, and saplings were more symptomatic than old growth trees. We suggest that these trends are related to differing sensitivity of the two species to O(3) exposure, and to higher O(3) exposures and drought stress that younger trees may have experienced during germination and establishment.     

Spatial and temporal small-scale variability of nitrogen mobilization in a forest ecosystem with high N deposition in NW-Germany

For conifer stands in NW-Germany with high DIN load (23-35 kg N ha⁻¹ a⁻¹) and a long history of nitrogen export the risk of N mobilization were investigated. Ammonium is the most mobilized N species, pointing towards either conditions not favoring nitrification or, more likely - under the dominant aerobic conditions - a very high amount of ammonium in the forest floor. Independence of net nitrification and net ammonification from each other indicates the existence of two separate systems. The nitrifying system depends very much on biotic conditions - as a function of energy and moisture - and seems not to be directly related to N deposition. In contrast, for the ammonification system (Oe horizon) a correlation with the sum of ammonium deposition three months prior to sampling was found. However, the role of disturbance, i.e. nitrogen export, during the last centuries and the role of recovery of the N balance during the last 150 years is still not clear.     

Evaluation of nitrogen dioxide photolysis rates in an urban area using data from the 1997 Southern California Ozone Study

The photolysis of nitrogen dioxide and formaldehyde are two of the most influential reactions in the formation of photochemical air pollution, and their rates are computed using actinic flux determined from a radiative transfer model. In this study, we compare predicted and measured nitrogen dioxide photolysis rate coefficients (j_NO2). We used the Tropospheric Ultraviolet-Visible (TUV) radiation transfer model to predict j_NO2 values corresponding to measurements performed in Riverside, California as part of the 1997 Southern California Ozone Study (SCOS’97). Spectrally resolved irradiance measured at the same site allowed us to determine atmospheric optical properties, such as aerosol optical depth and total ozone column, that are needed as inputs for the radiative transfer model. Matching measurements of aerosol optical depth, ozone column, and j_NO2 were obtained for 14 days during SCOS’97. By using collocated measurements of the light extinction caused by aerosols and ozone over the full height of the atmosphere as model input, it was possible to predict sudden changes in j_NO2 resulting from atmospheric variability. While the diurnal profile of the rate coefficient was readily reproduced, j_NO2 model predicted values were found to be consistently higher than measured values. The bias between measured and predicted values was 17–36%, depending on the assumed single scattering albedo. By statistical analysis, we restricted the most likely values of the single scattering albedo to a range that produced bias on the order of 20–25%. It is likely that measurement error is responsible for a significant part of the bias. The aerosol single scattering albedo was found to be a major source of uncertainty in radiative transfer model predictions. Our best estimate indicates its average value at UV-wavelengths for the period of interest is between 0.77 and 0.85.     

Acid deposition and the element composition of pine tree rings

The element contents of pine tree rings (Pinus sylvestris L.) were analyzed in five areas differing in the level of sulphur deposition. The elements studied and their average content in xylem were as follows:

aluminium 4.2, manganese 65, zinc 8.4, potassium 390, rubidium 2.2, calcium 930, magnesium 131, phosphorus 38, sulphur 56, nitrogen 0.65, boron 4.3 and molybdenum 0.03 mg/kg.

The contents of aluminium, potassium and rubidium were higher in tree rings from the 1950's on, particularly in southern Finland. The increase is probably due to acid deposition in the soil.     

Eutrophication trends in forest soils in Galicia (NW Spain) caused by the atmospheric deposition of nitrogen compounds

We calculated the sensitivity of Galician forest soils to eutrophication caused by atmospheric deposition of nitrogen compounds, using the Simple Mass Balance (SMB) method as described by [Posch, M., de Vries, W., Hettelingh, J.-P., 1995. Critical loads of sulphur and nitrogen. In: Posch, M., de Smet, P.A.M., Hettelingh, J.-P., Downing, R.J. Calculation and Mapping of Critical Thresholds in Europe. Status Report 1995, Coordination Center for Effects, National Institute for Public Health and the Environment, Bilthoven, The Netherlands, pp. 31-42]. Deposition values were used to calculate critical loads exceedance. Galician natural forest ecosystems can support nitrogen deposition loads of more than 10 kg Nha (-1) yr (-1). The lowest critical loads (approximately 10 kg Nha (-1) yr (-1)) mainly occurred in forest stands in the interior zone, while highest critical load values (approximately 68 kg Nha (-1) yr (-1)) were observed in eucalyptus stands at low altitudes in the littoral area. Exceedances based on N deposition levels, calculated from data recorded in 2001, occurred in 40% of the forest soils, showing the need to control N emissions in these areas to prevent possible eutrophication of soils and waters. Analysis of rainfall bulk composition revealed that ammonium, probably derived from agricultural and cattle activities, was the main compound responsible for N deposition in Galicia.     

Atmospheric nitrogen deposition in south-east Scotland: Quantification of the organic nitrogen fraction in wet,dry and bulk deposition

Water soluble organic nitrogen (WSON) compounds are ubiquitous in precipitation and in the planetary boundary layer, and therefore are a potential source of bioavailable reactive nitrogen. This paper examines weekly rain data over a period of 22 months from June 2005 to March 2007 collected in 2 types of rain collector (bulk deposition and “dry + wet” deposition) located in a semi-rural area 15 km southwest of Edinburgh, UK (N55°51′44″, W3°12′19″). Bulk deposition collectors are denoted in this paper as “standard rain gauges”, and they are the design used in the UK national network for monitoring precipitation composition. “Dry + wet” deposition collectors are flushing rain gauges and they are equipped with a rain detector (conductivity array), a spray nozzle, a 2-way valve and two independent bottles to collect funnel washings (dry deposition) and true wet deposition. On average, for the 27 weekly samples with 3 valid replicates for the 2 types of collectors, dissolved organic nitrogen (DON) represented 23% of the total dissolved nitrogen (TDN) in bulk deposition. Dry deposition of particles and gas on the funnel surface, rather than rain, contributed over half of all N-containing species (inorganic and organic). Some discrepancies were found between bulk rain gauges and flushing rain gauges, for deposition of both TDN and DON, suggesting biological conversion and loss of inorganic N in the flushing samplers.     

Regional and historical variation in the nitrogen content of Racomitrium lanuginosum in Britain in relation to atmospheric nitrogen deposition

The moss Racomitrium lanuginosum (Hedw.) Brid. is an important component of the drier parts of ombrotrophic mires and montane heaths in north-western Britain. The extent and quality of the montane heaths dominated by R. lanuginosum has declined in recent decades, perhaps in part due to the effects of acidic deposition at high elevations. This paper examines the effect of atmospheric nitrogen deposition, which has increased during this century, on the nitrogen content of R. lanuginosum in Britain. The nitrogen content of the moss reflects the magnitude of the atmospheric supply being least in north-western Scotland and greatest (as much as six-fold greater) near to urban centres in northern England. This regional difference was less marked (only approx. two-fold) during the 19th century (as revealed from the analysis of herbarium specimens) when nitrogen concentrations were appreciably lower. Transplant studies both between regions and between sites within a mountain system demonstrated the importance of atmospheric deposition in determining the tissue nitrogen concentration of the moss. The results are discussed in relation to the potential importance of the enhanced atmospheric nitrogen supply to the normally nitrogen-impoverished montane heaths, and to the growth and persistence of the moss.