Atmospheric deposition and soil water chemistry in Swedish forests since 1985 - Effects of reduced emissions of sulphur and nitrogen.

Trends for the atmospheric deposition of sulphur (S) and inorganic nitrogen (inorg-N) to forests and changes in the forest soil water chemistry in Sweden have been assessed since 1985, with special focus on the last 25 years, based on measurements within the Swedish Throughfall Monitoring Network (SWETHRO). The reductions in the deposition of S and inorg-N in the southern part of Sweden corresponded relatively well with the pollutant emission reductions for S and inorg-N from both EU27 + UK and Sweden during 1996/97-2021/22. For northern Sweden the deposition of S and inorg-N decreased to a lesser extent than both European and Swedish emissions. The bulk deposition of NO3-N has decreased more than the deposition of NH4-N over the last 25-year period, which is consistent with the much larger emission reductions for NOx compared to NH3 from EU27 + UK and Sweden. The S concentrations in the soil water, at 50 cm below soil surface, have decreased during the last 25 years, however somewhat less than the S deposition. At sites with low ANC and pH in the beginning of the period, the increase in ANC was generally greater and the increase in pH was smaller, but at sites with high pH and ANC above zero, the increase in pH was dominant, in line with the nonlinear relationship between pH and ANC in the soil water. The incidence of elevated concentrations of NO3-N in the soil water was highest in southwest Sweden, ranging between 4 and 19 % of all measuring occasions since 1985/86. The reduced deposition of N over the 35-year period was not reflected in the incidence of elevated concentrations of NO3-N in the soil water over time.

Twenty years of nitrogen deposition to Norway spruce forests in Sweden.

The yearly, total (dry+wet) deposition of inorganic nitrogen (inorg-N) to Norway spruce forests was estimated with a full spatial coverage over Sweden for a twenty-year period, 2001-2020, based on combined measurements with Teflon string samplers, throughfall deposition and bulk deposition to the open field. The results were based on a novel method to apply estimates of the dry deposition based on measurements at a limited number of sites, to a larger number of sites with only bulk deposition measurements, in turn based on the existence of a strong geographical gradient in the dry deposition of inorg-N from southwest to northeast Sweden. The method should be applicable for other geographical regions where gaseous NH3, NO2 and HNO3 are not main drivers of N dry deposition and where geographical gradients in dry deposition could be defined. It was shown that Norway spruce forests in south Sweden receive more N from deposition than has been previously estimated, based on modelling. Clear time trends were demonstrated for decreased deposition of inorg-N to Norway spruce forests in all parts of Sweden. The decreases were somewhat larger than what could be expected from the decrease in the reported emissions of inorg-N from Europe. The results emphasize that estimates of the total deposition are necessary in order to map levels and follow the development of N deposition in forests.

New flux based dose-response relationships for ozone for European forest tree species.

To derive O3 dose-response relationships (DRR) for five European forest trees species and broadleaf deciduous and needleleaf tree plant functional types (PFTs), phytotoxic O3 doses (PODy) were related to biomass reductions. PODy was calculated using a stomatal flux model with a range of cut-off thresholds (y) indicative of varying detoxification capacities. Linear regression analysis showed that DRR for PFT and individual tree species differed in their robustness. A simplified parameterisation of the flux model was tested and showed that for most non-Mediterranean tree species, this simplified model led to similarly robust DRR as compared to a species- and climate region-specific parameterisation. Experimentally induced soil water stress was not found to substantially reduce PODy, mainly due to the short duration of soil water stress periods. This study validates the stomatal O3 flux concept and represents a step forward in predicting O3 damage to forests in a spatially and temporally varying climate.

Estimates of AOT ozone indices from time-integrated ozone data and hourly air temperature measurements in southwest Sweden.

Surface ozone concentration and surface air temperature was measured hourly at three coastal sites, four low elevation inland sites and two high elevation inland sites in southwestern Sweden. Diurnal ozone concentration range (DOR) and diurnal temperature range (DTR) were strongly correlated, both spatially and temporally, most likely because both depended on atmospheric stability. Accumulated ozone exposure above a threshold concentration of x nmol mol(-1) (AOTx) was estimated from time-integrated ozone concentration (as from diffusive sampling) and measures of ozone concentration variability. Two methods both estimated 24-h AOTx with high accuracy (modelling efficiencies >90% for x

Increasing risk for negative ozone impacts on vegetation in northern Sweden.

Trends were found for increasing surface ozone concentrations during April-September in northern Sweden over the period 1990-2006 as well as for an earlier onset of vegetation growing season. The highest ozone concentrations in northern Sweden occurred in April and the ozone concentrations in April showed a strong increasing trend. A model simulation of ozone flux for Norway spruce indicated that the provisional ozone flux based critical level for forests in Europe is exceeded in northern Sweden. Future climate change would have counteracting effects on the stomatal conductance and needle ozone uptake, mediated on the one hand by direct effect of increasing air temperatures and on the other through increasing water vapour pressure difference between the needles and air. Thus, there is a substantial and increasing risk for negative impacts of ozone on vegetation in northern Sweden, related mainly to increasing ozone concentrations and an earlier onset of the growing season.

Promoting the O3 flux concept for European forest trees.

Tropospheric ozone (O3) levels are predicted to stay high, being a factor within "global change" with potential effects on the carbon sink strength of forest trees. Hence, new approaches to O3 risk assessment and their validation are required, although appropriate databases for adult trees are scant. Approaches based on external O3 exposure are presently being evaluated against the ones on O3 flux into leaves, as the cumulative uptake has the capacity for deriving O3 risk from cause-effect relationships. The effective dose, however, needs to account for the trees' O3 defence and tolerance in addition to O3 uptake. The current status of promoting the preferable mechanistic O3 flux concept is highlighted for major regions of Europe, addressing refinements and simplifications needed for routine use. At the pan-European scale, however, the flux-based concept is ready for use in O3 risk assessment and has the potential of meso-scale application at the forest ecosystem level.

Risk assessments for forest trees: the performance of the ozone flux versus the AOT concepts.

Published ozone exposure-response relationships from experimental studies with young trees performed at different sites across Europe were re-analysed in order to test the performance of ozone exposure indices based on AOTX (Accumulated exposure Over a Threshold of X nmol mol(-1)) and AF(st)Y (Accumulated Stomatal Flux above a threshold of Y nmol m(-2) s(-1)). AF(st)1.6 was superior, as compared to AOT40, for explaining biomass reductions, when ozone sensitive species with differing leaf morphology were included in the analysis, while this was not the case for less sensitive species. A re-analysis of data with young black cherry trees, subject to different irrigation regimes, indicated that leaf visible injuries were more strongly related to the estimated stomatal ozone uptake, as compared to the ozone concentration in the air. Experimental data with different clones of silver birch indicated that leaf thickness was also an important factor influencing the development of ozone induced leaf visible injury.

A cumulative ozone uptake-response relationship for the growth of Norway spruce saplings.

Norway spruce saplings [Picea abies (L.) Karst.] were exposed during four growing seasons to different ozone treatments in open-top chambers: charcoal filtered air (CF), non-filtered air (NF) and non-filtered air with extra ozone (NF+, 1.4xambient concentrations). The CF and NF+ ozone treatments were combined with phosphorous deficiency and drought stress treatments. The total biomass of the trees was harvested at different intervals during the experimental period. The ozone uptake to current-year needles of the Norway spruce saplings was estimated using a multiplicative stomatal conductance simulation model. There was a highly significant correlation between the reduction of total biomass and the estimated cumulative ozone uptake, which did not vary when different thresholds were applied for the rate of ozone uptake. The reduction of the total biomass was estimated to 1% per 10 mmol m(-2) cumulated ozone uptake, on a projected needle area basis.

Impact of ozone on the growth of birch (Betula pendula) saplings.

Saplings of one half-sib family of birch, Betula pendula, were exposed to three levels of ozone in open-top chambers (OTCs) during two growing seasons 1997-1998. The ozone treatments were non-filtered air (NF, accumulated daylight AOT40 over the two growing seasons of 3.0 l l-1 h), non-filtered air with extra ozone (NF+, accumulated daylight AOT40 of 27.3 l l-1 h) and non-filtered air with additional extra ozone (NF++, accumulated daylight AOT40 of 120 l l-1 h). The birch saplings, including the roots, were harvested after the first and second growing seasons. After the first growing season, the NF++ treatment reduced the total wood biomass by 22%, relative to the NF treatment. There was no further reduction of the total wood biomass in the NF++ treatment after the second growing season. The root biomass was reduced by 30% after the first growing season. The shoot/root ratio, as well as the proportional biomass of leaves, were increased by ozone during both years. The ozone impact on the relative growth rate was estimated to -2% per 10 l l-1 h daylight AOT40 per growing season.

Impact of ozone and reduced water supply on the biomass accumulation of Norway spruce saplings.

Norway spruce saplings [Picea abies (L.) Karst.] were exposed during four growing seasons to two different ozone treatments in open-top chambers: charcoal filtered air (CF), and non-filtered air with extra ozone (NF+, 1.4xambient concentrations). Within each ozone treatment the saplings were either kept well watered or treated with a 7-8 week period with reduced water supply each growing season. The total biomass of the trees was measured in April and September during each of the last three growing seasons. NF+ significantly reduced the total biomass accumulation of Norway spruce saplings during the fourth growing season. No interaction between ozone and reduced water supply could be detected. The magnitude of the ozone impact after 4 years of exposure was an 8% reduction of the total plant biomass and a 1.5% reduction of the RGR. The reduced water supply reduced the total biomass 29% and the RGR 12%.

Simulations of stomatal conductance and ozone uptake to Norway spruce saplings in open-top chambers.

A simulation model was developed to estimate the stomatal conductance and ozone flux to Norway spruce saplings in open-top chambers. The model was parameterized against needle conductance measurements that were made on 4-6-year-old spruce saplings, grown in open-top chambers, in July-September during three different seasons. The spruce saplings were either maintained well watered or subject to a 7-8 week drought period in July-September each year. The simulated conductance showed a good agreement with the measured conductance for the well-watered as well as the drought stress-treated saplings. The simulations were significantly improved when different vapour pressure deficit (VPD) functions were applied for well-watered and drought-stressed spruce saplings. The cumulated ozone uptake which was calculated from the conductance simulations showed less variation between years, compared to the cumulative ozone exposure index AOT40 (accumulated exposure over a threshold of 40 ppb or nl l(-1)) for the corresponding time periods. Measurements in May 1995 demonstrated the occurrence of long-term 'memory-effects' from the drought stress treatments on the conductance. Memory-effects need to be considered when simulation models for stomatal conductance are to be applied to long-lived forest trees under a multiple stress situation.

An ozone flux-response relationship for wheat.

Six open-top chamber experiments with field-grown wheat Triticum aestivum L. (five with spring wheat and one with winter wheat) were combined to test which of the two ozone exposure indices, AOT40 and CFO(3), that provided the most consistent relationship between relative yield loss and ozone exposure. AOT40 is the accumulated exposure over a threshold ozone concentration of 40 nl l(-1), while CFO(3) is the cumulative flux of ozone (uptake) to the flag leaves. The ozone uptake of the flag leaves was estimated using a stomatal conductance model, sensitive to phenology, light, vapour pressure deficit (VPD) and temperature in combination with measurements of the boundary layer conductance in the open-top chambers. Both indices were calculated for the grain-filling period, defined as the time from anthesis until 2 weeks before harvest. The duration of the grain-filling period was shown to be closely related to the rate of accumulation of thermal time above a base temperature of 0 degrees C. The CFO(3) index provided a much more consistent pattern in terms of ozone effects compared to the AOT40 index. This was especially the case for spring wheat, for which a linear regression between relative yield and CFO(3) using all five data sets is presented. According to the stomatal conductance model, VPD limited daytime stomatal conductance in warm and dry years, while temperature was the most important limiting factor during daytime in cool and humid years. The effect of light was mainly to delimit the time period of the day during which substantial uptake of ozone took place. It is concluded that, compared to the AOT40 index, the more mechanistically relevant flux-based index CFO(3) will estimate larger yield loss in the relatively humid parts of western and northern Europe, while smaller yield loss will be estimated for the dry summer climates in south and central Europe. The use of an ozone flux threshold, similar to the cut-off concentration 40 nl l(-1) in AOT40, did not improve the performance of the CFO(3) index.

Rapid and Specific Modulation of Stomatal Conductance by Blue Light in Ivy (Hedera helix) : An Approach to Assess the Stomatal Limitation of Carbon Assimilation.

Low intensity (0.015 millimole per square meter per second) blue light applied to leaves of Hedera helix under a high intensity red light background (0.50 millimole per square meter per second red light) induced a specific stomatal opening response, with rapid kinetics comparable to those previously reported for stomata with ;grass type' morphology. The response of stomatal conductance to blue light showed a transient ;overshoot' behavior at high vapor pressure difference (2.25 +/- 0.15 kiloPascals), but not at low vapor pressure difference (VPD) (0.90 +/- 0.10 kilo-Pascal). The blue light-induced conductance increase was accompanied by an increase in net photosynthetic carbon assimilation, mediated by an increase in the intercellular concentration of carbon dioxide. Values of assimilation once the blue light-stimulated conductance increase reached steady state were less than those at the peak of the overshoot, but the ratios of assimilation to transpiration (A/E) and blue light-stimulated DeltaA/DeltaE were greater during the steady-state response than during the overshoot. These results indicate that significant stomatal limitation of assimilation can occur, but that this limitation may improve water use efficiency under high VPD conditions. Under high intensity red light, the decline in A/E associated with an increase in VPD was minimized when conductance was stimulated by additional low intensity blue light. This effect indicates that the blue light response of stomata may be important in H. helix for the optimization of water use efficiency under natural conditions of high irradiance and VPD.