A unifying explanation for variation in ozone sensitivity among woody plants.

Tropospheric ozone is considered the most detrimental air pollutant for vegetation at the global scale, with negative consequences for both provisioning and climate regulating ecosystem services. In spite of recent developments in ozone exposure metrics, from a concentration-based to a more physiologically relevant stomatal flux-based index, large-scale ozone risk assessment is still complicated by a large and unexplained variation in ozone sensitivity among tree species. Here, we explored whether the variation in ozone sensitivity among woody species can be linked to interspecific variation in leaf morphology. We found that ozone tolerance at the leaf level was closely linked to leaf dry mass per unit leaf area (LMA) and that whole-tree biomass reductions were more strongly related to stomatal flux per unit leaf mass (r2  = 0.56) than to stomatal flux per unit leaf area (r2  = 0.42). Furthermore, the interspecific variation in slopes of ozone flux-response relationships was considerably lower when expressed on a leaf mass basis (coefficient of variation, CV = 36%) than when expressed on a leaf area basis (CV = 66%), and relationships for broadleaf and needle-leaf species converged when using the mass-based index. These results show that much of the variation in ozone sensitivity among woody plants can be explained by interspecific variation in LMA and that large-scale ozone impact assessment could be greatly improved by considering this well-known and easily measured leaf trait.

Epidemiological analysis of ozone and nitrogen impacts on vegetation - Critical evaluation and recommendations.

For human health studies, epidemiology has been established as important tool to examine factors that affect the frequency and distribution of disease, injury, and other health-related events in a defined population, serving the purpose of establishing prevention and control programs. On the other hand, gradient studies have a long tradition in the research of air pollution effects on plants. While there is no principal difference between gradient and epidemiological studies, the former address more one-dimensional transects while the latter focus more on populations and include more experience in making quantitative predictions, in dealing with confounding factors and in taking into account the complex interplay of different factors acting at different levels. Epidemiological analyses may disentangle and quantify the contributions of different predictor variables to an overall effect, e.g. plant growth, and may generate hypotheses deserving further study in experiments. Therefore, their use in ecosystem research is encouraged. This article provides a number of recommendations on: (1) spatial and temporal aspects in preparing predictor maps of nitrogen deposition, ozone exposure and meteorological covariates; (2) extent of a dataset required for an analysis; (3) choice of the appropriate regression model and conditions to be satisfied by the data; (4) selection of the relevant explanatory variables; (5) treatment of interactions and confounding factors; and (6) assessment of model validity.

Past, present and future concentrations of ground-level ozone and potential impacts on ecosystems and human health in northern Europe.

This review summarizes new information on the current status of ground-level ozone in Europe north of the Alps. There has been a re-distribution in the hourly ozone concentrations in northern Europe during 1990-2015. The highest concentrations during summer daytime hours have decreased while the summer night-time and winter day- and night-time concentrations have increased. The yearly maximum 8-h mean concentrations ([O3]8h,max), a metric used to assess ozone impacts on human health, have decreased significantly during 1990-2015 at four out of eight studied sites in Fennoscandia and northern UK. Also the annual number of days when the yearly [O3]8h,max exceeded the EU Environmental Quality Standard (EQS) target value of 60ppb has decreased. In contrast, the number of days per year when the yearly [O3]8h,max exceeded 35ppb has increased significantly at two sites, while it decreased at one far northern site. [O3]8h,max is predicted not to exceed 60ppb in northern UK and Fennoscandia after 2020. However, the WHO EQS target value of 50ppb will still be exceeded. The AOT40 May-July and AOT40 April-September metrics, used for the protection of vegetation, have decreased significantly at three and four sites, respectively. The EQS for the protection of forests, AOT40 April-September 5000ppbh, is projected to no longer be exceeded for most of northern Europe sometime before the time period 2040-2059. However, if the EQS is based on Phytotoxic Ozone Dose (POD), POD1, it may still be exceeded by 2050. The increasing trend for low and medium range ozone concentrations in combination with a decrease in high concentrations indicate that a new control strategy, with a larger geographical scale than Europe and including methane, is needed for ozone abatement in northern Europe.

Biomass burning in eastern Europe during spring 2006 caused high deposition of ammonium in northern Fennoscandia.

High air concentrations of ammonium were detected at low and high altitude sites in Sweden, Finland and Norway during the spring 2006, coinciding with polluted air from biomass burning in eastern Europe passing over central and northern Fennoscandia. Unusually high values for throughfall deposition of ammonium were detected at one low altitude site and several high altitude sites in north Sweden. The occurrence of the high ammonium in throughfall differed between the summer months 2006, most likely related to the timing of precipitation events. The ammonia dry deposition may have contributed to unusual visible injuries on the tree vegetation in northern Fennoscandia that occurred during 2006, in combination with high ozone concentrations. It is concluded that long-range transport of ammonium from large-scale biomass burning may contribute substantially to the nitrogen load at northern latitudes.

Acidification trends in south Swedish forest soils 1986-2008 - slow recovery and high sensitivity to sea-salt episodes.

Soil water chemistry in forest soils over 20 years was studied at nine sites in southern Sweden. The aim was to investigate the recovery from acidification and the influence of strong sea salt episodes that occur in the region. All sites but one showed signs of recovery from acidification along with the reduced sulphur deposition, but the recovery progress was slow and the soil water was in most cases still highly acidic at the end of the period. In several cases the recovery was delayed by episodes of sea salt deposition, leading to transient acidification. The less marked decrease of sulphur concentrations in soil water than of sulphur deposition, highlighted the importance of sulphur adsorption/desorption in the acidification and recovery process. Nitrogen retention capacity was exceeded on one site, leading to nitrate leaching and extremely low pH. Storm fellings on two sites in the end of the period led to effects similar to those of regeneration fellings. It was concluded that the soils in the region are in an early stage of recovery. The future progress of recovery strongly depends on future nitrogen retention of forest soils and the frequency of sea salt episodes.

Reduced European emissions of S and N--effects on air concentrations, deposition and soil water chemistry in Swedish forests.

Changes in sulphur and nitrogen pollution in Swedish forests have been assessed in relation to European emission reductions, based on measurements in the Swedish Throughfall Monitoring Network. Measurements were analysed over 20 years with a focus on the 12-year period 1996 to 2008. Air concentrations of SO(2) and NO(2), have decreased. The SO(4)-deposition has decreased in parallel with the European emission reductions. Soil water SO(4)-concentrations have decreased at most sites but the pH, ANC and inorganic Al-concentrations indicated acidification recovery only at some of the sites. No changes in the bulk deposition of inorganic nitrogen could be demonstrated. Elevated NO(3)-concentrations in the soil water occurred at irregular occasions at some southern sites. Despite considerable air pollution emission reductions in Europe, acidification recovery in Swedish forests soils is slow. Nitrogen deposition to Swedish forests continues at elevated levels that may lead to leaching of nitrate to surface waters.

Assessing the risk of N leaching from forest soils across a steep N deposition gradient in Sweden.

Nitrogen leaching from boreal and temporal forests, where normally most of the nitrogen is retained, has the potential to increase acidification of soil and water and eutrophication of the Baltic Sea. In parts of Sweden, where the nitrogen deposition has been intermediate to high during recent decades, there are indications that the soils are close to nitrogen saturation. In this study, four different approaches were used to assess the risk of nitrogen leaching from forest soils in different parts of Sweden. Nitrate concentrations in soil water and C:N ratios in the humus layer where interpreted, together with model results from mass balance calculations and detailed dynamic modelling. All four approaches pointed at a risk of nitrogen leaching from forest soils in southern Sweden. However, there was a substantial variation on a local scale. Basing the assessment on four different approaches makes the assessment robust.

Ozone exposure and impacts on vegetation in the Nordic and Baltic countries.

Ozone concentrations are generally considerably lower over northern Europe as compared with continental and southern Europe. However, ozone becomes toxic for vegetation mainly after it has been taken up into the leaf interior through the stomata. The rates of ozone uptake into the leaves are, somewhat simplified, the product of the air ozone concentrations and the degree of stomatal opening. The phytotoxic impacts of ozone can be almost as important in northern Europe as they are in continental and southern Europe. The long daylight hours as well as the rather humid environment conditions, both in the air and soil, promote stomatal openings in northern Europe. This article summarizes scientific evidence that supports the conclusion that ozone abatement policies regarding vegetation in Europe, as well as in the rest of the world, should be based on estimates of the leaf ozone uptake and not only on the ozone concentration in the air.

Evidence for impacts of near-ambient ozone concentrations on vegetation in southern Sweden.

Substantial impacts of near-ambient ozone concentrations on agricultural crops, trees, and seminatural vegetation are demonstrated for southern Sweden. Impacts of ambient ozone levels (2-15 microL L(-1) hr annual accumulated ozone exposure over a threshold of 40 nL L(-1) [AOT40]) range from a 2%-10% reduction for trees (e.g., leaf chlorophyll, tree growth) up to a 15% reduction for crops (e.g., yield, wheat/potato). Visible leaf injury on bioindicator plants caused by ambient ozone levels has been clearly demonstrated. The humid climatic conditions in Sweden promote high rates of leaf ozone uptake at a certain ozone concentration. This likely explains the comparatively large ozone impacts found for vegetation in southern Sweden at relatively low ozone concentrations in the air. It is important that the future methods used for the representation of ozone impacts on vegetation across Europe are based on the leaf ozone uptake concept and not on concentration-based exposure indices, such as AOT40.

Synoptic weather types and long-range transport patterns for ozone precursors during high-ozone events in southern Sweden.

We studied long-range transport patterns and related weather types in relation to high-ozone events in southern Sweden. The aim was to deepen the understanding of the relationship between Lamb-Jenkinson weather types and surface ozone concentration variation, thus widening the application of the weather type analysis of air quality at 4 sites in this region. The long-range transport patterns associated with high-ozone events were classified into trajectories from Western Europe, Eastern Europe, and in the vicinity of southern Sweden (VIC). The VIC type, characterized by short and whirling curves, represented more than 40% of the high-ozone events at the studied rural sites. More than half of the high-ozone events occurred under high-pressure conditions, belonging to weather type A (anticyclones). The high correlation coefficient between annual counts of weather type A and those of long-range transport pattern VIC confirmed the strong link between stagnant weather conditions and high-ozone events, especially during the summer. Furthermore, a strong linear anticorrelation was detected between high-ozone events and annual counts of weather type C (cyclones) during the summer. This relationship implies that the frequency of weather type C is a useful indicator for low risk of summertime high-ozone events in southern Sweden. Moreover, the relationship between the weather type and high ozone risk may be useful in examining the potential effect of climate change on the regional air quality.

Ozone impairs autumnal resorption of nitrogen from birch (Betula pendula) leaves, causing an increase in whole-tree nitrogen loss through litter fall.

Saplings of one half-sib family of birch, Betula pendula Roth, were exposed to three ozone concentrations (non-filtered air (NF); non-filtered air + 10-20 nmol O(3) mol(-1) (NF+); non-filtered air + 40-60 nmol O(3) mol(-1) (NF++)) in open-top chambers during two growing seasons from 1997 to 1998. Shed leaves were collected regularly during both growing seasons and, in 1998, the dry mass (DM) and nitrogen (N) concentrations ([N]) of the shed leaves were measured to quantify the total amount of N lost through litter fall. Dry mass and [N] were also determined in mid-August for attached, mature and non-senescent leaves, in order to estimate autumnal leaf N resorption efficiency and proportional leaf DM decrease. Net photosynthetic capacity was measured during August and September 1998, in a population of leaves that emerged in mid-July. Photosynthesis declined with increasing leaf age in the NF++ treatment, whereas it remained high throughout the measurement period in the NF and NF+ treatments. In both years, leaves abscised prematurely in the NF++ treatment, whereas this effect was only significant in 1998 in the NF+ treatment. There was a strong linear relationship between proportional leaf shedding and daylight ozone exposure above a threshold of 40 nmol mol(-1) (daylight AOT40) during the growing season. The resorption of N was significantly impaired by ozone, and the smaller autumnal decrease in leaf DM in elevated ozone concentrations suggested that the bulk resorption of leaf DM was also inhibited. Nitrogen resorption efficiencies were 81, 73 and 63% and leaf mass decreases were 45, 36 and 30% in the NF, NF+ and NF++ treatments, respectively. Compared with the NF treatment, total N loss through litter fall was increased by 16 and 122% in the NF+ and NF++ treatments, respectively. We conclude that ozone impaired N resorption from birch leaves before abscission, causing a substantial increase in whole-tree N loss through litter fall.

Economic assessment of the negative impacts of ozone on crop yields and forest production. A case study of the estate Ostads Säteri in southwestern Sweden.

Ground level ozone concentrations, in combination with the prevailing climate, at the estate Ostads Säteri in southwestern Sweden were estimated to reduce the yield of wheat and potato ranging between 5% and 10%. Occasionally, in years with the highest ozone concentrations and/or climatic conditions favoring high rates of ozone uptake to the leaves, yield loss levels above 10% may occur. Based on simple extrapolation, these ozone-induced reductions of crop yields at Ostads Säteri represent a potential total annual yield loss in Sweden in the range of 24.5 million Euro for wheat and 7.3 million Euro for potato, respectively. A simulation of forest growth at Ostad Säteri predicted that prevailing mean ozone exposure during 1993-2003 had the potential to reduce forest growth by 2.2% and the economic return of forest production by 2.6%. Using this value for extrapolation to the national level, the potential annual economic loss for Sweden due to negative impacts of ozone on forest production would be in the range of 56 million Euro (2004 prices).

Cell structural changes in the needles of Norway spruce exposed to long-term ozone and drought.

Effects of ozone and/or drought on Norway spruce needles were studied using light microscopy and electron microscopy. Saplings were exposed to ozone in open-top chambers during 1992-1995 and also to drought in the late summers of 1993-1995. Samples from current and previous year needles were collected five times during 1995. Ozone increased the numbers of peroxisomes and mitochondria, which suggests that defence mechanisms against oxidative stress were active. The results from peroxisomes suggest that the oxidative stress was more pronounced in the upper side of the needles, and those from mitochondria that defence was more active in the younger needle generation. Possibly due to the good nitrogen status and the active defence, no ozone-specific chloroplast alterations were seen. At the end of the season, older needles from ozone treatments had smaller central vacuoles compared with other needles. Cytoplasmic vacuoles around the nucleus were increased by ozone in the beginning of the experiment, and did not increase towards the end of the season as in the controls. These results from vacuoles may indicate that ozone affected the osmotic properties of the cells. Decreased number and underdevelopment of sclerenchyma cells and proliferation of tonoplast were related to nutrient imbalance, which was enhanced by drought. Larger vascular cylinders and more effective starch accumulation before and after the drought periods compensated for the reduced water status. Numbers of peroxisomes and mitochondria were increased in the drought-exposed needles before the onset of drought treatments of the study year, i.e. these changes were memory effects. Interactions between ozone and drought were few.

Clover as a tool for bioindication of phytotoxic ozone--5 years of experience from southern Sweden--consequences for the short-term critical levels.

Critical levels (CLs) for ozone effects on plants in Europe have been defined within the UN-ECE Convention on Long-Range Transboundary Air Pollution, CLRTAP. The purpose of the short-term CLs is to ensure protection of all crops to acute ozone injury. The currently used CLs are based on the ozone exposure of the plants during daylight hours expressed as AOT40 (Accumulated exposure Over the Threshold 40 nmol mol(-1) ozone). The aims of this study were: to test the performance of the current short-term CLs, to test alternative ozone exposure indices and to test if changes in the ozone cut-off concentration, the inclusion of a lag-period (LP) between exposure and identification of visible ozone injury or the duration of the ozone integration period improved the performance of the exposure index. The analysis was based on 38 different datasets from experiments with subterranean clover, Trifolium subterraneum in southern Sweden. AOT indices generally performed better than averaged ozone concentrations or SUM (Sum of ozone concentrations when a threshold is exceeded) indices. Regression analysis showed that the current short-term CL, AOT40 with a VPD (water vapour pressure deficit) threshold of 1.5 kPa, explained 56% of the variation in visible injury. A longer exposure period and the introduction of a LP, admitting visible ozone injury time to develop after exposure, improved the performance of the exposure index. AOT30 accumulated over 10 days before harvest, excluding a LP of 3 days before injury observation, performed best and explained 88% of the variation in visible injury. AOT40 indices left a rather large amount of visible injury unexplained indicating that a lower cut-off concentration for ozone is preferable. The results of the investigation indicated that a visible injury threshold of 10% improved the distinction between harmful and less harmful exposure.

Impact of four years exposure to different levels of ozone, phosphorus and drought on chlorophyll, mineral nutrients, and stem volume of Norway spruce, Picea abies.

Saplings of one clone of Norway spruce, Picea abies (L.) Karst, were planted in 120 l pots in 1991 and exposed to three levels of ozone, two levels of phosphorus and two levels of water supply in 42 open-top chambers (OTCs), during 1992-1996. The effects of pots and OTCs were also tested. Nutrient concentrations of the needles were not affected by ozone, while the low phosphorus supply (LP) and drought stress (D) treatments had significant effects on several mineral nutrients, e.g. phosphorus, calcium, magnesium, manganese, sulphur and boron. Ozone reduced the chlorophyll concentration in the 2- and 3-year-old needles in 1994 and 1995. The highest ozone concentration reduced the stem volumes (- 8%), as well as the stem lengths (- 5%), of the saplings in 1993 and 1994, after two and three years of exposure. After the fourth growing season this ozone-induced reduction in stem volume disappeared which might be caused by pot limitation. LP supply and D both caused large decreases in the stem volume and length. The needles from LP treatment had as high P concentration as 1.2-1.5 mg g-1, implying a need for increasing the critical value for phosphorus. The OTC enclosure stimulated the stem volume growth significantly compared to saplings growing in ambient plots. This was suggested to be attributed to the slightly higher temperature in the OTCs. The overall result is that ozone in southern Sweden is likely to have negative effects on Norway spruce trees, although much less than other environmental factors, e.g. water and phosphorus.