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PubMed1992

Metabolic pathways in Paracoccus denitrificans and closely related bacteria in relation to the phylogeny of prokaryotes.

Denitrification and methylotrophy in Paracoccus denitrificans are discussed. The properties of the enzymes of denitrification: the nitrate-nitrite antiporter, nitrate reductase, nitrite reductase, nitric oxide reductase and nitrous oxide reductase are described. The genes for none of these proteins have yet been cloned and sequenced from P. denitrificans. A number of sequences are available for enzymes from Escherichia coli, Pseudomonas stutzeri and Pseudomonas aeruginosa. It is concluded that pathway specific c-type cytochromes are involved in denitrification. At least 40 genes are involved in denitrification. In methanol oxidation at least 20 genes are involved. In this case too pathway specific c-type cytochromes are involved. The sequence homology between the quinoproteins methanol dehydrogenase, alcoholde-hydrogenase and glucose dehydrogenase is discussed. This superfamily of proteins is believed to be derived from a common ancestor. The moxFJGI operon determines the structural components of methanol dehydrogenase and the associated c-type cytochrome. Upstream of this operon 3 regulatory proteins were found. The moxY protein shows the general features of a sensor protein and the moxX protein those of a regulatory protein. Thus a two component regulatory system is involved in both denitrification and methylotrophy. The phylogeny of prokaryotes based on 16S rRNA sequence is discussed. It is remarkable that the 16S rRNA of Thiosphaera pantotropha is identical to that of P. denitrificans. Still these bacteria show a number of differences. T. pantotropha is able to denitrify under aerobic circumstances and it shows heterotrophic nitrification. Nitrification and heterotrophic nitrification are found in species belonging to the beta-and gamma-subdivisions of purple non-sulfur bacteria. Thus the occurrence of heterotrophic nitrification in T. pantotropha, which belongs to the alpha-subdivision of purple non-sulfur bacteria is a remarkable property. Furthermore T. pantotropha contains two nitrate reductases of which the periplasmic one is supposed to be involved in aerobic denitrification. The nitrite reductase is of the Cu-type and not of the cytochrome cd1 type as in P. denitrificans. Also the cytochrome b of the Qbc complex of T. pantotropha is highly similar to its counterpart in P. denitrificans. It is hypothesized that the differences between these two organisms which both contain large megaplasmids is due to a combination of loss of genetic information and plasmid-coded properties. The distribution of a number of complex metabolic systems in eubacteria and in a number of species belonging to the alpha-group of purple non sulphur bacteria is reviewed.(ABSTRACT TRUNCATED AT 400 WORDS)

Bacteria↗
PubMed1993

The effects of atmospheric nitrogen deposition on the soil chemistry of coniferous forests in The Netherlands.

Nitrogen fluxes, particularly those of ammonium, are extremely high in Dutch forests. In soils exposed to high ammonium deposition, acidification, eutrophication or a combination of both processes may occur. In addition to the amounts of ammonium deposited, the rate of soil nitrification determines which process takes place. A nation-wide investigation, in which three coniferous tree species were involved, was carried out to study the relation between deposition fluxes, measured by means of throughfall and bulk samplers, and the chemical composition of the soil. The ammonium deposition accounted directly for the high ammonium content and the high ammonium/cation ratios in the soil. In the top layer of most of the forest soils which were investigated nitrification rates were low. In these stands ammonium/cation ratios in the soil often reflected ammonium/cation ratios in throughfall water. Even in soils with relatively high nitrification rates, ammonium concentrations exceeded those of nitrate in the top layer of the mineral soil, indicating that ammonium deposition was more important than nitrification rate in determining the predominant form of nitrogen.

Journal Article↗
PubMed2006

Modelling of biological processes during aerobic treatment of piggery wastewater aiming at process optimisation.

A dynamic mathematical model was developed for the simulation of the aerobic treatment of piggery wastewater. This model includes the carbon oxidation, the nitrification and the denitrification. According to the experimental results obtained during this study, a modified version of the activated sludge model No. 1 has been developed. The model includes (1) nitrite as intermediate of nitrification and denitrification, (2) the distinction between the anoxic heterotrophic yield and the aerobic heterotrophic yield, respectively equal to 0.53 and 0.6 and (3) the first-order hydrolysis of the slowly biodegradable fraction. The calibration and the validation of the model was performed using experimental data from three experiments with two piggery wastewaters. A set of kinetic and stoichiometric parameters emerged from these tests. Except the kinetic of hydrolysis of the slowly biodegradable organic matter varying from 6 to 25 gCOD(gCODday)(-1), all other parameters were similar for all experiments. The dissolved oxygen concentration was identified as the main variable influencing the nitrite accumulation during nitrification. In the calibrated model, the oxygen half-saturation coefficient of the ammonium oxidisers (0.3g O(2)m(-3)) was lower than for the nitrite oxidisers (1.1 gO(2)m(-3)), leading to nitrite accumulation when the dissolved oxygen concentration was low. Simulations with the proposed model could be very useful for improved design and management of biological treatment of piggery wastewaters, particularly in case of partial nitrification to nitrite directly followed by denitrification.

Aerobiosis↗
PubMed2005

Effects of increased deposition of atmospheric nitrogen on an upland Calluna moor: N and P transformations.

This study determined the effects of increased N deposition on rates of N and P transformations in an upland moor. The litter layer and the surface of the organic Oh horizon were taken from plots that had received long-term additions of ammonium nitrate at rates of 40, 80 and 120 kg N ha(-1) yr(-1). Net mineralisation processes were measured in both field and laboratory incubations. Soil phosphomonoesterase (PME) activity and rates of N(2)O release were measured in laboratory incubations and root-surface PME activity measured in laboratory microcosms using Calluna vulgaris bioassay seedlings. Net mineralisation rates were relatively slow, with net ammonification consistently stimulated by N addition. Net nitrification was marginally stimulated by N addition in the laboratory incubation. N additions also increased soil and root-surface (PME) activity and rates of N(2)O release. Linear correlations were found between litter C:N ratio and all the above processes except net nitrification in field incubations. When compared with data from a survey of European forest sites, values of litter C:N ratio were greater than a threshold below which substantial, N input-related increases in net nitrification rates occurred. The maintenance of high C:N ratios with negligible rates of net nitrification was associated with the common presence of ericaceous litter and a mor humus layer in both this moorland as well as the forest sites.

Air Pollutants↗
PubMed2005

Modelling of nitrogen release from MBT waste.

The "LaNDy" model (landfill nitrogen dynamics model) is a new mathematical tool for the evaluation of the long-term behaviour of nitrogen in mechanical-biologically pretreated (MBP) waste. LaNDy combines a hydraulic model based on RICHARD's equation with one-dimensional heat flow in landfills, kinetics of biological degradation, gas diffusion, nitrification and denitrification. A suitable temperature-dependent N mineralisation sub-model was based on numerous data from the literature and own LSR-experiments. With the "nitrification modus" of the LaNDy model, kinetic data of nitrification, thermodynamic data of denitrification and diffusion characteristics of gaseous components (especially of oxygen and methane) are used as an additional input for the preliminary calculation of the long-time impact of nitrification and denitrification. Examples of predicted temperature distribution and leachate ammonium concentrations, using different landfill size, age of the landfill (10 to approximately 100 a) and hydraulic conductivity of the MBP waste, are presented in this paper.

Forecasting↗
PubMed2006

A quantitative measure of nitrifying bacterial growth.

Nitrifying bacteria convert ammonia (NH3) to nitrate (NO3-) in a nitrification reaction. Methods to quantitatively separate the growth rate of these important bacterial populations from that of the dominant heterotrophic bacteria are important to our understanding of the nitrification process. The changing concentration of ammonia is often used as an indirect measure of nitrification but ammonification processes generate ammonia and confound this approach while heterotrophs remove nitrate via denitrification. Molecular probe methods can tell us what proportion of the microbial community is nitrifying bacteria but not their growth rate. The technique proposed here was able to quantify the growth rate of the nitrifying bacterial populations amidst complex ecological processes. The method incubates [methyl-3H] thymidine with water samples in the presence and absence of an inhibitor of nitrification-thiourea. The radioactively labeled DNA in the growing bacteria was extracted. The rate of incorporation of the label into the dividing bacterial DNA was used to determine bacterial growth rate. Total bacterial community growth rates in full-scale and pilot-scale fixed-film nitrifying reactors and an activated sludge reactor were 2.1 x 10(8), 4.1 x 10(8) and 0.4 x 10(8)cell ml(-1)d(-1), respectively; the growth rate of autotrophic-nitrifying bacteria was 0.7 x 10(8), 2.6 x 10(8) and 0.01 x 10(8)cell ml(-1)d(-1), respectively. Autotrophic-nitrifying bacteria contributed 30% and 60% of the total bacterial community growth rate in the nitrifying reactors whereas only 2% was observed in the activated sludge reactor that was not designed to nitrify. The rates of ammonia loss from the nitrifying reactors corresponded to the rate of growth of the nitrifying bacteria. This method has the potential to more often identify factors that enhance or limit nitrifying processes in both engineered and natural aquatic environments.

Bacteria↗
PubMed2002

Nitrifying and heterotrophic population dynamics in biofilm reactors: effects of hydraulic retention time and the presence of organic carbon.

Two biofilm reactors operated with hydraulic retention times of 0.8 and 5.0 h were used to study the links between population dynamics and reactor operation performance during a shift in process operation from pure nitrification to combined nitrification and organic carbon removal. The ammonium and the organic carbon loads were identical for both reactors. The composition and dynamics of the microbial consortia were quantified by fluorescence in situ hybridization (FISH) with rRNA-targeted oligonucleotide probes combined with confocal laser scanning microscopy, and digital image analysis. In contrast to past research, after addition of acetate as organic carbon nitrification performance decreased more drastically in the reactor with longer hydraulic retention time. FISH analysis showed that this effect was caused by the unexpected formation of a heterotrophic microorganism layer on top of the nitrifying biofilm that limited nitrifiers oxygen supply. Our results demonstrate that extension of the hydraulic retention time might be insufficient to improve combined nitrification and organic carbon removal in biofilm reactors.

Acetates↗
PubMed2002

Simultaneous P and N removal in a sequencing batch biofilm reactor: insights from reactor- and microscale investigations.

A sequencing batch biofilm reactor (SBBR) with well established enhanced biological phosphate removal (EBPR) was subjected to higher ammonium concentrations to stimulate and eventually implement simultaneous nitrification. Changes of activity and populations were investigated by a combination of online monitoring, microsensor measurements and fluorescence in situ hybridisation (FISH) of biofilm sections. Nitrification and nitrifying bacteria were always restricted to the periodically oxic biofilm surface. Both, activity and population size increased significantly with higher ammonium concentrations. Nitrification always showed a delay after the onset of aeration, most likely due to competition for oxygen by coexisting P accumulating and other heterotrophic bacteria during the initial aeration phase. This view is also supported by comparing oxygen penetration and oxygen uptake rates under low and high ammonium conditions. Therefore, simultaneous nitrification and phosphorus removal in a P removing SBBR appears to be only possible with a sufficiently long oxic period to ensure oxygen availability for nitrifiers.

Bacteria↗
PubMed2003

Evaluation of the impact of bioaugmentation and biostimulation by in situ hybridization and microelectrode.

Three rotating disk biofilm reactors were operated to evaluate whether bioaugmentation and biostimulation can be used to improve the start-up of microbial nitrification. The first reactor was bioaugmented during start-up period with an enrichment culture of nitrifying bacteria, the second reactor received a synthetic medium containing NH(4)(+) and NO(2)(-) to facilitate concomitant proliferation of ammonia- and nitrite-oxidizing bacteria, and the third reactor was used as a control. To evaluate the effectiveness of bioaugmentation and biostimulation approaches, time-dependent developments of nitrifying bacterial community and in situ nitrifying activity in biofilms were monitored by fluorescence in situ hybridization (FISH) technique and microelectrode measurements of NH(4)(+), NO(2)(-), NO(3)(-), and O(2). In situ hybridization results revealed that addition of the enrichment culture of nitrifying bacteria significantly facilitated development of dense nitrifying bacterial populations in the biofilm shortly after, which led to a rapid start-up and enhancement of in situ nitrification activity. The inoculated bacteria could proliferate and/or survive in the biofilm. In addition, the addition of nitrifying bacteria increased the abundance of nitrifying bacteria in the surface of the biofilm, resulting in the higher nitrification rate. On the other hand, the addition of 2.1mM NO(2)(-) did not stimulate the growth of nitrite-oxidizing bacteria and did inhibit the proliferation of ammonia-oxidizing bacteria instead. Thus, the start-up of NO(2)(-) oxidation was unchanged, and the start-up of NH(4)(+) oxidation was delayed. In all the three biofilm reactors, data sets of time series analyses on population dynamics of nitrifying bacteria determined by FISH, in situ nitrifying activities determined by microelectrode measurements, and the reactor performances revealed an approximate agreement between the appearance of nitrifying bacteria and the initiation of nitrification activity, suggesting that the combination of these techniques was a very powerful monitoring tool to evaluate the effectiveness of bioaugmentation and biostimulation strategies.

Biofilms↗
PubMed2003

Coupled BAS and anoxic USB system to remove urea and formaldehyde from wastewater.

Wastewater containing formaldehyde and urea was treated using a coupled system consisting of a biofilm airlift suspension (BAS) reactor and an anoxic upflow sludge blanket (USB) reactor. The anoxic USB reactor was used to carry out denitrification and urea hydrolysis, while the BAS reactor was used to carry out nitrification. In a first step, individual experiments were carried out to investigate the effects of both compounds on the nitrifying and denitrifying biomass. The BAS reactor was fed with a synthetic medium containing 500 mg N-NH4(+)l(-1) and 100mg N-urea l(-1), that were added continuously to this medium. Neither urea hydrolysis nor inhibition of nitrification was observed. Nitrification efficiency decreased when formaldehyde was fed during shocks at concentrations of 40, 80 and 120 mg C-formaldehyde l(-1). The anoxic USB reactor was fed with a synthetic medium containing nitrate, formaldehyde and urea. Concentrations of formaldehyde in the reactor of 100-120 mg C-formaldehyde l(-1) caused a decrease in the denitrification and urea hydrolysis rates. In a second step, the coupled system was operated at recycling ratios (R) of 3 and 9. Fed C/N ratios of 0.58, 1.0 and 1.5 g C-formaldehyde g(-1) N-NH4(+) were used for every recycling ratio. The maximum nitrogen removal percentages were achieved at a C/N ratio of 1.0 g C-formaldehyde g(-1) N-NH4(+) for both recycling ratios. A fed C/N ratio of 1.5 g C-formaldehyde g(-1) N-NH4(+) caused a decrease in the efficiency of the system with respect to nitrogen removal, due to the presence of formaldehyde in the BAS reactor, which decreased the nitrification. Formaldehyde was completely removed in the BAS reactor and a heterotrophic layer formed around the nitrifying biofilm.

Biofilms↗
PubMed2003

Characterization and evaluation of aerobic granules in sequencing batch reactor.

In order to investigate the aerobic granules cultured under alternating aerobic and anoxic conditions, a sequencing batch reactor (SBR) was operated without the presence of a carrier material. Nitrification and denitrification occurred alternately in the SBR operation, with an increased nitrification efficiency of up to 97% and a high chemical oxygen demand (COD) removal efficiency of up to 95%. It was observed that physical characteristics of granule play an important role in the performance of the SBR process. Light microscopy was used to observe the time dependent development of the granules in the SBR. Based on the microscopic observations, some floc-like sludges remained in the form of a mixture with granules for 30 days of operation. Even though various granule sizes had been formed in the reactor after 50 days, the granule sizes were primarily from 1 +/- 0.35 to 1.3 +/- 0.45 mm, rarely exceeding 2 mm. The granules were analyzed by a combination of microelectrodes and fluorescent in situ hybridization (FISH), which provides more detailed information on what happens inside the granules. Based on their results, ammonia oxidizing bacteria (AOB) existed primarily in the upper and middle layers of the granule. Assuming a first-order reaction for nitrification, most of the nitrification is likely to occur from the surface to 300 microm into the granular thickness.

Bacteria, Aerobic↗
PubMed2001

Performance evaluation of single-sludge reactor system treating high-strength nitrogen wastewater.

In the single-sludge reactor system treating high-strength nitrogen wastewater (similar to anaerobically pretreated piggery wastewater), the NH4(+)-N removal efficiencies (98-82%) are higher than total nitrogen removal efficiencies (71-43%). The mixed liquor recycle ratio only imposes a slight effect on total nitrogen removal efficiency. The alkalinity change data could be used for monitoring and control of the reactor system. To evaluate the performance of the single-sludge reactor system, a simplified nitrification-denitrification model (with nitrification capacity, denitrification capacity, and denitrification potential concepts) and a graphically analytical technique are proposed. It turns out that ammonia nitrification and total nitrogen removal efficiencies are strongly dependent on the process load and reactor configuration, and an optimal operating condition requires a proper match between nitrification and denitrification.

Aerobiosis↗
PubMed2003

Influence of hydraulic loading and air flowrate on urban wastewater nitrogen removal with a submerged fixed-film reactor.

Nutrient disposal to sensitive areas, particularly nitrogen and phosphorus from wastewater treatment plants, provokes eutrophication reducing water quality. Fixed film technology is widely used for the removal of organic matter and nitrogen by the biological process of nitrification-denitrification. This paper studies a nitrification and post-denitrification lab-scale plant with a downflow aerobic submerged filter for removal of organic matter and nitrification, followed by an anoxic upflow biofilter for denitrification. Recycled construction material (clay shists) was employed as support material and methanol was used as carbon source. After 2 weeks of acclimation in which nitrification reached steady-state conditions, different hydraulic loadings (0.35-1.59 m(3)/m(2)h) and air flowrates (7.78-43.5 m(3)/m(2)h) were applied for 1 year. The highest hydraulic loading which complied with the EU regulation on nitrogen disposal was 0.71 m(3)/m(2)h (1.6 h). Hydraulic retention time (HRT), which corresponded to a nitrogen removal of 0.64 kg N/m(3) per day operating at an air flowrate of 25.6 m(3)/m(2)h. Concerning to organic matter removal efficiency, the aerobic reactor accepted a maximum chemical oxygen demand (COD) volumetric loading of 16.0 kg COD/m(3) per day with a 75% COD removal efficiency. For all the tests carried out, suspended solids (SS) concentration in the outlet water was less than 35 mg/l.

Bacteria, Aerobic↗
PubMed2003

Accelerated degradation of methyl iodide by agrochemicals.

The fumigant methyl iodide (MeI, iodomethane) is considered a promising alternative to methyl bromide (MeBr) for soil-borne pest control in high-cash-value crops. However, the high vapor pressure of MeI results in emissions of a significant proportion of the applied mass into the ambient air, and this may lead to pollution of the environment. Integrating the application of certain agrochemicals with soil fumigation provides a novel approach to reduce excessive fumigant emissions. This study investigated the potential for several agrochemicals that are commonly used in farming operations, including fertilizers and nitrification inhibitors, to transform MeI in aqueous solution. The pseudo-first-order hydrolysis half-life (t(1/2)) of MeI was approximately 108 d, while the transformation of MeI in aqueous solutions containing selected agrochemicals was more rapid, with t(1/2) < 100 d (t(1/2) < 0.5 d in some solutions containing nitrification inhibitors). The influence of these agrochemicals on the rate of MeI degradation in soil was also determined. Adsorption to soil apparently reduced the availability of some nitrification inhibitors in the soil aqueous phase and lowered the degradation rate in soil. In contrast, addition of the nitrification inhibitors thiourea and allylthiourea to soil significantly accelerated the degradation of MeI, possibly due to soil surface catalysis. The t(1/2) of MeI was <20 h in thiourea- and allylthiourea-amended soil, considerably less than that in unamended soil (t(1/2) > 300 h).

Adsorption↗
PubMed2003

Dicyandiamide and 3,4-dimethyl pyrazole phosphate decrease N2O emissions from grassland but dicyandiamide produces deleterious effects in clover.

The application of nitrogen fertilisers leads to different ecological problems such as nitrate leaching and the release of nitrogenous gases. N2O is a gas involved in global warming, therefore, agricultural soils can be regarded as a source of global warming. Soil N2O production comes from both the nitrification and denitrification processes. From an ecological viewpoint, using nitrification inhibitors with ammonium based fertilisers may be a potential management strategy to lower the fluxes of N2O, thus decreasing its undesirable effect. In this study, the nitrification inhibitors (NIs) dicyandiamide (DCD) and 3,4-dimethyl pyrazole phosphate (DMPP) have been evaluated as management tools to mitigate N2O emissions from mineral fertilisation and slurry application in grassland systems (experiments 1 and 2), and to assess the phytotoxic effect of these inhibitors per se on clover (experiment 3). Both nitrification inhibitors acted in maintaining soil nitrogen (N) in ammonium form, decreasing cumulative N2O emissions. DCD, but not DMPP, produced phytotoxic effects and yield reduction in white clover. A nutrient imbalance, which led to a senescence process visually observed as chlorosis and necrosis at the border of the leaves, was noted.

Fertilizers↗
PubMed2006

Carbon and nitrogen removal in a granular bed baffled reactor.

The application of an anaerobic five compartment granular bed baffled reactor (GRABBR) was investigated with brewery wastewater for combined carbon and nitrate removal, with a separate downstream nitrification unit for converting ammonia to nitrate. The GRABBR was operated at an organic loading rate of 3.57 kg chemical oxygen demand (COD) m(-3) d(-1) and ammoniacal nitrogen (NH4-N) loading rate of 0.13 kg NH4-N m(-3) d(-1) when nitrified effluent from a downstream nitrification unit was recycled to the feed point of the GRABBR. Carbonaceous matter and nitrate were removed simultaneously in the GRABBR at different recycle to influent ratios (from 1 to 2), with nitrogen oxide (nitrate and nitrite nitrogen, NOx-N) loading rates varying from 0.04 to 0.05 kg NOx-N m(-3) d(-1). At all recycle to influent ratios, COD removal efficiency of 97% to 98% were observed in the GRABBR, and over 99% by the two-stage treatment configuration (i.e. GRABBR and nitrification unit). All the nitrates added to the GRABBR were denitrified in the first three compartments of the system. For all the recycle to influent ratios studied, almost all ammonia was converted to nitrate nitrogen with only small traces of nitrite nitrogen in the nitrification unit. Methane production was observed throughout the experimental period with its composition varying from 25% to 50%, showing that simultaneous methanogenesis and denitrification occurred. This study shows that a GRABBR could bring about a high degree of carbon and nitrate removal, with simultaneous methanogenesis and denitrification, due to plug flow granular bed multi-stage characteristics of the bioreactor.

Ammonia↗
PubMed2005

Microbial characteristics and nitrogen transformation in planted soil filter for domestic wastewater treatment.

We studied an experimental horizontal subsurface-flow planted sand filter in Kodijärve, Estonia. We measured the microbial biomass, nitrogen immobilization, potential nitrification, soil respiration, multiple carbon source utilization patterns of the microbial consortia of the soil samples, the carbon, nitrogen, and phosphorus content of the soil samples, the water quality and physicochemical indicators in water sampling wells as well as emissions of CO2, N2, NO2, and CH4 from the two beds (the dry bed and the wet bed) in the wetland. The potential nitrification of the upper layer of the dry bed could not be attributed primarily to autotrophic nitrification, or the nitrifying bacteria in this layer could be facultative heterotrophs, whereas autotrophic nitrification is predominant in the upper layer of the wet bed. It also was found that changing aeration conditions in the lower layer of the dry bed have resulted in a lower diversity of the microbial community and led to a relative depletion of easily degradable soil carbon resources.

Biomass↗
PubMed2001

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.

California↗