Uptake of SO2 in shoots of Scots pine: field measurements of net flux of sulphur in relation to stomatal conductance

Abstract. Net fluxes of gaseous sulphur compounds, water vapour and carbon dioxide to current- and one-year-old shoots of Scots pine were simultaneously measured in the field. The shoots were fumigated in temperature-controlled assimilation chambers with sulphur dioxide at concentrations ranging from 40 to 250 μg m⁻³ (15–95 ppb). The hypothesis that stomatal opening regulates the uptake of sulphur dioxide was tested. The following conclusions were reached concerning the dry deposition of sulphur dioxide to a dry coniferous shoot.
There was a marked diurnal variation in the uptake rate of sulphur dioxide. The net deposition velocity of sulphur was about three times higher during the day than during the night during July-October. Stomatal opening was not the primary factor controlling the dry deposition rate of sulphur dioxide. The net dry deposition rate was significantly smaller than the calculated rate based on stomatal conductances for water vapour.
Part of this deviation could be explained by re-emission of reduced sulphur compound(s) from the needles. The re-emission of sulphur was a light-dependent process and might be of great significance for the sulphur balance calculations.
A variable deposition velocity should be used in atmospheric transport models to account for the diurnal and seasonal variation in the surface resistance of a dry canopy.     

The simulation of soil organic matter and nitrogen accumulation in Scots pine plantations on bare parent material using the combined forest model EFIMOD

The individual-based combined forest model EFIMOD including the soil-sub model SOMM has been used for the simulation of Scots pine stand growth and soil organic matter (SOM) accumulation on a humus-free bare mineral surface. The growth of Scots pine plantation, with an initial density of 10 000 trees ha⁻¹ and average tree biomass of 0.01 kg was simulated for 50 yr under Central European climatic conditions (i) with varying atmospheric nitrogen inputs and (ii) different rates of initial application of raw undecomposed organic material or compost, on humus-free parent material. The accumulation of typical raw humus was simulated in all cases. The accumulation was most intensive in the simulation of high atmospheric nitrogen input. The humus pool in the mineral topsoil was small but achieved its maximum value with compost application. SOM nitrogen accumulation was scant in all cases, except the compost applications with low atmospheric nitrogen input. No statistically significant differences of SOM and stand parameters were found between variants without organic matter and those with low input of organic manure. However, the maximum relative rate of SOM and nitrogen accumulation was found in the scenario without organic manure, under slowly growing unstable Scots pine plantation. This revised version was published online in June 2006 with corrections to the Cover Date. This revised version was published online in June 2006 with corrections to the Cover Date. This revised version was published online in June 2006 with corrections to the Cover Date.     

Atmospheric deposition and watershed nitrogen export along an elevational gradient in the Catskill Mountains, New York

Cumulative effects of atmospheric N deposition mayincrease N export from watersheds and contribute tothe acidification of surface waters, but naturalfactors (such as forest productivity and soildrainage) that affect forest N cycling can alsocontrol watershed N export. To identify factors thatare related to stream-water export of N, elevationalgradients in atmospheric deposition and naturalprocesses were evaluated in a steep, first-orderwatershed in the Catskill Mountains of New York, from1991 to 1994.Atmospheric deposition of SO ₄ ^2– , andprobably N, increased with increasing elevation withinthis watershed. Stream-water concentrations ofSO ₄ ^2– increased with increasing elevationthroughout the year, whereas stream-waterconcentrations of NO ₃ ^– decreased withincreasing elevation during the winter and springsnowmelt period, and showed no relation with elevationduring the growing season or the fall. Annual exportof N in stream water for the overall watershed equaled12% to 17% of the total atmospheric input on thebasis of two methods of estimation. This percentagedecreased with increasing elevation, from about 25%in the lowest subwatershed to 7% in the highestsubwatershed; a probable result of an upslope increasein the thickness of the surface organic horizon,attributable to an elevational gradient in temperaturethat slows decomposition rates at upper elevations. Balsam fir stands, more prevalent at upper elevationsthan lower elevations, may also affect the gradient ofsubwatershed N export by altering nitrification ratesin the soil. Variations in climate and vegetationmust be considered to determine how future trends inatmospheric deposition will effect watershed export ofnitrogen.     

Effect of atmospheric ammonia on the nitrogen metabolism of Scots pine (Pinus sylvestris) needles

Four-year-old seedlings of Scots pine ( Pinus sylvestris L.) were exposed to filtered air (FA), and to FA supplemented with NH₃ (60 and 240 μg m⁻³) in controlled-environment chambers for 14 weeks. Exposure to the higher NH₃ concentration resulted in an increased activity of glutamine synthetase (GS, EC 6.3.1.2), and an increase in the concentrations of soluble proteins, total nitrogen, free amino acids and leaf pigments in the needles. The GS activity (μmol g⁻¹ fresh weight h⁻¹) in the needle extract increased to levels 69% higher than in FA and the soluble protein concentration to levels 22% higher. Total nitrogen concentration in the needles was 42% higher than in FA, while the free amino acid concentration was 300% higher, which was caused by an increase in arginine, glutamate, aspartate and glutamine. Chlorophyll a , chlorophyll b and carotenoid concentrations were 29, 38 and 11% higher, respectively. Neither the glutamate dehydrogenase (GDH, EC 1.4.1.2) activity nor the concentrations of free NH₄⁺ and glucose in the needles were affected by exposure to NH₃. After NH₃ fumigation at 240 μg m⁻³ the starch concentration decreased by 39% relative to the FA. The results indicate that the metabolism of Scots pine acclimates to concentrations of NH₃ which are 3 to 10 times higher than the average concentration in areas with intensive stock farming. The possible mechanisms underlying acclimation to NH₃ are discussed.     

Effect of ectomycorrhizal fungi on nitrogen mineralisation and the growth of Scots pine seedlings in natural peat

The aim of this study was to investigate the potential of different isolates of ectomycorrhizal (EM) fungi to enhance the growth of Pinus sylvestris seedlings in five natural peat substrates with different nitrogen concentrations, and the effect of the Scots pine seedlings and fungal inoculum on the formation of dissolved inorganic and organic nitrogen in peat. Utilization of different organic nitrogen compounds by microbial community in the peat was also investigated using Biolog MT MicroPlates. Inoculation of the seedlings with EM fungi enhanced seedling growth. Piloderma croceum increased root growth especially, whereas Lactarius rufus increased needle growth and Suillus variegatus I, II and III improved both root and needle growth. All the EM fungi also significantly affected stem growth. Nitrogen concentration of the peat did not affect seedling growth as much as the EM fungi. At the lowest peat N concentration (1.17%) NH ₄ ⁺ mineralisation was lower and DON (dissolved organic nitrogen) accumulation higher than at higher peat N concentrations. The EM fungal isolates had different effects on NH ₄ ⁺ and DON accumulation/degradation in peat. The EM fungal isolates significantly increased NH ₄ ⁺ formation in the peat, whereas L. rufus and P. croceum had an opposite effect on DON accumulation. S. variegatus I significantly decreased the DON concentrations during peat incubation. The N concentration of the peat slightly affected the utilization of amino acids and polyamines by the microbial community, whereas inoculation with S. variegatus I, II or III had no effect.     

Light microscopic structure of needles of Scots pine (Pinus sylvestris L.) in relation to air pollution and needle element concentrations in S.E. Finland and the Karelian Isthmus, N.W. Russia

 The mesophyll structure of current and second year needles of mature Scots pines (Pinus sylvestris L.), growing on a survey line from the Karelian Isthmus (N.W. Leningrad region, Russia) to S.E. Finland, was studied at the light microscopic level in 1992 and 1994. The area is affected by sulphur dioxide and nitrogen oxide emissions from Russia (200  000–285 000 ton/year SO₂ and about 100  000 ton/year NO_X), Estonia (140  000–320  000 ton/year SO₂ and 30  000–50  000 ton/year NO_X) and Finland (6000–9000 ton/year SO₂ and 23 000–25 000 ton/year NO_X) and by unknown amounts of neutralizing compounds from Estonia. Mean concentrations of SO₂ were between 1.0 and 1.5 μg m^–3 and those of ozone between 52 and 61 μg m^–3 during growing seasons of 1991–1994. The concentrations tended to be lower in 1993/94 than in 1991/92. The results showed cellular changes related to exposure to sulphur dioxide, ozone and acid wet deposition. The percentage of cells with injuries related to acid deposition was higher in 1994 than in 1992, possibly due to higher reduction of neutralizing than of acidifying compounds in the main emission areas, while the percentage of cells with injuries related to sulphur dioxide correlated positively with needle sulphur concentrations and showed a declining trend between Karelian Isthmus and S.E. Finland in the needles sampled in 1992. These changes and those related to ozone were considerably less numerous in 1994 than in 1992, possibly on account of the lower concentrations of SO₂ and ozone in 1993/94 than in 1991/92, especially in early summer, the time of the most sensitive phase of needle growth. In addition, periods of lower wet deposition in summer 1992, and considerably lower temperatures in autumn/winter 1992 than in 1994 may have enhanced the injuries in the needles collected in October/December 1992, relative to those collected in 1994. The results of the microscopic work agreed well with the simultaneous chemical measurements, pointing to the usefulness of cellular analysis for diagnostic work in the field. Received: 26 September 1997 / Accepted: 24 November 1997     

Assimilation of nitrate and ammonium by the Scots pine (Pinus sylvestris) seedling under conditions of high nitrogen supply

Seedlings of Scots pine ( Pinus sylvestris L.) were grown on perlite for 21 days under controlled conditions. Apart from the water control, KNO₃ (15 m M ), (NH₄)₂SO₄ (7.5 m M ), and NH₄NO₃ (15 m M ) were offered to study the effects of a high nitrogen supply on nitrogen assimilation. In some experiments 1.3 m M potassium was added to the basic ammonium solutions. In labelling studies nitrate and ammonium were 2.3 atom%¹⁵N-enriched. It was found that over the 21-day period approximately three times more ammonium-N was taken up than nitrate-N. However, nitrate and ammonium, applied simultaneously, were taken up to the same extent as if they were applied separately (additivity). The presence of K⁺ in the medium did not affect N-uptake. Among the soluble N-containing compounds nitrate, ammonium and 8 amino acids were quantified. It was found that assimilation of nitrate can cope with the uptake of NO⁻₃ under all circumstances. Neither free nitrate nor ammonium or amino acids accumulated to an extent exceeding the values of water-grown seedlings. On the other hand, in case of high ammonium supply considerably more nitrogen was taken up than could be incorporated into nonsoluble N-containing substance ('protein'). The remaining nitrogen was found to accumulate in intermediary storage pools (free NH₄⁺, glutamine, asparagine, arginine). Part of this accumulated N could be incorporated into protein when potassium was offered in the nutrient solution. It is concluded that potassium is a requirement for a high rate of protein synthesis not only in crop plants but also in conifers.     

模拟氮沉降对温带典型森林土壤有效氮形态和含量的影响*

通过室内模拟氮沉降试验,研究了氮沉降对温带典型森林土壤有效氮的影响.结果表明:试验期间,与对照相比,经过氮沉降处理的土壤铵态氮、硝态氮和有效氮均呈增长的趋势,增加的程度取决于森林类型、土层、氮处理类型和氮处理的持续时间.氮沉降对不同林型土壤有效氮形态和含量的影响不同,氮沉降对混交林的影响弱于阔叶林,强于针叶人工纯林;土壤A层对氮沉降的敏感程度大于土壤B层;铵态氮形态沉降对土壤铵态氮含量的影响比对土壤硝态氮含量的影响大,而硝态氮形态沉降对土壤硝态氮含量的影响比对土壤铵态氮含量的影响大,混合形态的氮沉降对二者均有促进作用,且增加幅度更高;氮沉降对土壤有效氮的影响存在累加效应.     

Persistent effects of low concentrations of SO2 and NO2 on photosynthesis in Scots pine (Pinus sylvestris) needles

In an open-field experiment, 50-year-old trees of Scots pine (Pinus sylvestris L.) were fumigated with low concentrations of SO₂ and NO₂ (10–15 nl I^?1) during the growing season in four consecutive years (1988 to 1991). Results from the autumn and early winter of 1991 and 1992 are presented. The maximum photochemical efficiency of photosystem II (PSII), as indicated by the ratio of variable to maximum fluorescence (F_v/F_M) was assessed in current and one-year-old needles from the top and the bottom of the canopy. Furthermore, simultaneous measurements of photosynthetic O₂ evolution and chlorophyll fluorescence were made in current-year needles at 20°C. In general, the F_v/F_M ratio as well as the gross rate of O₂ evolution in needles of fumigated trees was not significantly different from that in needles of control trees during the fumigation period. However, both current and one-year-old needles sampled in November and December 1991 from the top of the canopy of fumigated trees had significantly lower F_v/F_M values than corresponding needles of control trees. Similar differences in F_v/F_M correlated with the treatments were observed in needles from the bottom of the canopy, indicating that the depression of F_v/F_M in needles of fumigated trees was not due to an increased susceptibility to photoinhibition. In 1992, when no fumigation occurred, differences in F_v/F_M between the treatments were not significant during autumn and early winter. The gross rate of O₂ evolution at high irradiances was significantly lower in current-year needles of fumigated trees sampled in November and December 1991 than in those of control trees. Furthermore, a nearly identical linear relationship between the quantum yield of PSII electron transport determined from chlorophyll fluorescence and the quantum yield of O₂ evolution (gross rate of O₂ evolution/PPFD) was found during autumn and early winter. This appeared to be largely a result of changes in the thermal energy dissipation within PSII. The observed differences in photosynthetic characteristics correlated with the different treatments after the fumigation period is suggested to be mainly caused by increased sensitivity of the needles of fumigated trees to low and subfreezing temperatures. However, current-year needles of fumigated trees tended to have a lower N content than those of control trees, which may partly explain the differences in gross photosynthesis between fumigated and control trees.     

The effects of SO2 and O3 fumigation on acid deposition and foliar leaching in the Liphook forest fumigation experiment

The ambient pollution climate at the Liphook forest fumigation site, where coniferous trees were fumigated with SO₂ and O₃, for 4 years under field conditions, was characteristic of the fringes of the areas where pollutant effects are a problem. Experimental treatments increased SO₂ concentrations to levels more characteristic of Eastern Europe, and summer O₃ concentrations by 30%. Deposition of SO₂ to the soil between the trees (inferred from shallow lysimeters) was significant, the deposition velocity being 2–1 mms^?1. Deposition to Scots pine and Sitka spruce canopies was greater, deposition velocities being 8.5 and 9.4 mm s^?1, respectively. These high values may perhaps be explained by co-deposition with NH₃. Calculations assume that dry deposition was the sole source of SO₄^2? gain in throughfall, and that there was no significant retention by the trees. There was a trend for O₃ to enhance SO₂ deposition to both soil and trees. Fumigation with SO₂ led to a significant increase in leaching of cations from foliage. Each species neutralized about 63% of the dry-deposited SO₂, predominantly by ion exchange for Ca and K. Equations are provided which allow calculation of foliar leaching given SO₂ concentrations or SO₄^2? deposition. Fumigation increased the rate of nutrient cycling considerably, without affecting foliar concentrations or damaging the trees. Ozone treatments did not enhance foliar leaching, calling into question some suggested mechanisms for the causes of forest decline.     

Interactions of elevated CO2, NH3 and O3 on mycorrhizal infection,gas exchange and N metabolism in saplings of Scots pine

Four-year-old saplings of Scots pine (Pinus sylvestris) (L.) were exposed for 11 weeks in controlled-environment chambers to charcoad-filtered air, or to charcoal-filtered air supplemented with NH₃ (40 g m^–3), O₃ (110 g m^–3 during day/ 40 g m^–3 during night) or NH₃+O₃. All treatments were carried out at ambient (259 L L^–1) and at elevated CO₂ concentration (700 L L^–1). Total tree biomass, mycorrhizal infection, net CO₂ assimilation (P_n), stomatal conductance (g_s), transpiration of the shoots and NH₃ metabolization of the needles were measured. In ambient CO₂ (1) gaseous NH₃ decreased mycorrhizal infection, without significantly affecting tree biomass or N concentration and it enhanced the activity of glutamine synthetase (GS) and glutamate dehydrogenase (GDH) in one-year-old needles; (2) ozone decreased mycorrhizal infection and the acitivity of GS in the needles, while it increased the activity og GDH; (3) exposure to NH₃+O₃ lessened the effects of single exposures to NH₃ and O₃ on reduction of mycorrhizal infection and on increase in GDH activity. Similar lessing effects on mycorrhizal infection as observed in trees exposed to NH₃+O₃ at ambient CO₂, were measured in trees exposed to NH₃+O₃ at elevated CO₂. Exposure to elevated CO₂ without pollutants did not significantly affect any of the parameters studied, except for a decrease in the concentration of soluble proteins in the needles. Elevated CO₂ _NH₃ strongly decreased root branching and mycorrhizal infection and temporarily stimulated P_n and g_s. The exposure to elevated CO₂+NH₃+O₃ also transiently stimulated P_n. The possible mechanisms underlying and integrating these effects are discussed. Elevated CO₂ clearly did not alleviate the negative effects of NH₃ and O₃ mycoorhiral infection. The significant reduction of mycorrhizal infection after exposure to NH₃ or O₃, observed before significant changes in gas exchange or growth occurred, suggest the use of mycorrhizal infection as an early indicator for NH₃ and O₃ induced stress.Abbreviations DW dry weight - FA filtered air - FA_a filtered air at ambient CO₂ - FA_e filtered air at elevated CO₂ - FW fresh weight - GDH glutamate dehydrogenase - GS glutamine synthetase - g_s stomatal conductance - P_n net CO₂ assimilation - RWR root weight ratio - SRL specific root length     

Use of stable isotopes to quantify nitrogen, potassium and magnesium dynamics in young Scots pine (Pinus sylvestris)

Two-yr-old Scots pine ( Pinus sylvestris ) seedlings were grown in sand culture for 1 yr with a generous supply of a balanced nutrient solution. Trees were repotted into clean sand in February 1998 and given either a reduced or adequate nutrient supply containing enriched ¹⁵N, ⁴¹K and ²⁶Mg to label nutrient uptake during spring 1998. Trees doubled their biomass during the experiment. Whole-tree net photosynthesis was reduced by 43% after 95 d in trees that received the lower nutrient supply ( P < 0.001), although differences in biomass between the two treatments were less pronounced. Remobilization contributed 83, 82 and 52% of the N, K and Mg, respectively, used to support growth of new tissues in trees that received reduced nutrient supply. Those receiving the higher nutrient supply still obtained 44–59% of nutrients used for spring growth of new tissues from remobilization. Current nutrient supply had no significant effect on the amount of N or Mg remobilized to new tissues but K remobilization was less in trees that received the lower nutrient supply ( P = 0.025). The importance of remobilization in young trees and problems associated with quantifying internal cycling of nutrients are discussed.     

Atmospheric nitrogen deposition has caused nitrogen enrichment and eutrophication of lakes in the northern hemisphere

We compiled chemical data and phytoplankton biomass (PB) data (chlorophyll a ) from unproductive lakes in 42 different regions in Europe and North America, and compared these data to inorganic nitrogen (N) deposition over these regions. We demonstrate that increased deposition of inorganic N over large areas of Europe and North America has caused elevated concentrations of inorganic N in lakes. In addition, the unproductive lakes in high N deposition areas had clearly higher PB relative to the total phosphorus (P) concentrations illustrating that the elevated inorganic N concentrations has resulted in eutrophication and increased biomass of phytoplankton. The eutrophication caused by inorganic N deposition indicates that PB yield in a majority of lakes in the northern hemisphere is (was) limited by N in their natural state. We, therefore, suggest that P limitation largely concerns lakes where the balance between N and P has been changed because of increased anthropogenic input of N.     

Ammonium,nitrate and dissolved organic nitrogen in seepage water as affected by compost amendment to European beech,Norway spruce,and Scots pine forests

Fertilization of nutrient-depleted and degraded forest soils may be required to sustain utilization of forests. In some European countries, the application of composts may now be an alternative to the application of inorganic fertilizers because commercial compost production has increased and compost quality has been improved. There is, however, concern that compost amendments may cause increased leaching of nitrogen, trace metals and toxic organic compounds to groundwater. The objective of this study was to assess the risk of ammonium (NH₄ ⁺), nitrate (NO₃ ^–) and dissolved organic nitrogen (DON) leaching following a single compost application to silty and sandy soils in mature beech (Fagus sylvatica L.), pine (Pinus silvestris L.) and spruce (Picea abies Karst.) forests at Solling and Unterlüß in Lower Saxony, Germany. Mature compost from separately collected organic household waste was applied to the soil surface at a rate of 6.3 kg m^–2 in the summer of 1997 and changes in NH₄ ⁺, NO₃ ^– and DON concentrations in throughfall and soil water at 10 and 100 cm soil depths were determined for 32 months. The spruce forests had the highest N inputs by throughfall water and the highest N outputs in both the control and compost plots compared with the pine and beech forests. Overall, the differences in total N outputs at 100 cm soil depth between the control and compost plots ranged between 0.3 and 11.2 g N m^–2 for the entire 32-month period. The major leaching of these amounts occurred during the first 17 months after compost amendments, but there was no significant difference in total N outputs (–0.2 to 1.8 g N m^–2) between the control and compost plots during the remaining 15 months. Most of the mineral soils acted as a significant sink for NO₃ ^– and DON as shown by a reduction of their outputs from 10 to 100 cm depth. Based on these results, we conclude that application of mature compost with high inorganic N contents could diminish the groundwater quality in the first months after the amendments. A partial, moderate application of mature compost with low inorganic N content to nutrient depleted forest soils can minimize the risk of NO₃ ^– leaching.     

Significance of litter- and humus-layer quality for rates and forms of N cycling in moder- to rawhumus-moder profiles under Scots pine (Pinus sylvestris L.)

Plant and Soil - In three Scots pine stands in northeastern Germany along a gradient of air pollutant deposition loads (particularly sulfate and alkaline dust), the net N release by decomposition,...     

Changes in species richness and composition in European acidic grasslands over the past 70 years: the contribution of cumulative atmospheric nitrogen deposition

Our study investigates the negative impact of nitrogen (N) deposition on species richness in acidic grasslands, based on a temporal comparison of vegetation data spanning a period of almost 70 years. We compiled a large data base of plots assigned to the Violion caninae grassland type, composed of managed, but unfertilized semi-natural grasslands on nutrient-poor, acidic soils. In total 1114 plots, mainly from Great Britain, the Netherlands and Germany, were compiled, dating back to 1939. Environmental site information included geographical and soil (mean Ellenberg values) variables as well as estimates of cumulative N and sulphur (S) deposition since 1939. Statistical analyses were carried out separately for the data subsets from the three regions. In all regions, the vegetation differentiation was mainly related to soil acidity and nutrient availability, as well as to the year of sampling and the cumulative amounts of N and S deposition. Plot-species richness of vascular plants and bryophytes (analysed for Great Britain only) decreased with time and analyses suggest these are affected by various factors, notably soil pH, but also latitude and cumulative N deposition. The latter explained more of the variation in species number than the year of sampling and cumulative S deposition, which supports the interpretation that the decline in species richness is mainly caused by increasing N availability and less by altered management and soil acidification. For Great Britain and Germany, cumulative N deposition showed a strong negative relationship with several biodiversity measures, especially the proportion of dicots, whereas it was positively related to the proportion of grass species. In general, our results give temporal evidence for the negative effect of N deposition on species richness in semi-natural vegetation.