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.     

Modelling stomatal ozone flux across Europe

A model has been developed to estimate stomatal ozone flux across Europe for a number of important species. An initial application of this model is illustrated for two species, wheat and beech. The model calculates ozone flux using European Monitoring and Evaluation Programme (EMEP) model ozone concentrations in combination with estimates of the atmospheric, boundary layer and stomatal resistances to ozone transfer. The model simulates the effect of phenology, irradiance, temperature, vapour pressure deficit and soil moisture deficit on stomatal conductance. These species-specific microclimatic parameters are derived from meteorological data provided by the Norwegian Meteorological Institute (DNMI), together with detailed land-use and soil type maps assembled at the Stockholm Environment Institute (SEI). Modelled fluxes are presented as mean monthly flux maps and compared with maps describing equivalent values of AOT40 (accumulated exposure over threshold of 40 ppb or nl l(-1)), highlighting the spatial differences between these two indices. In many cases high ozone fluxes were modelled in association with only moderate AOT40 values. The factors most important in limiting ozone uptake under the model assumptions were vapour pressure deficit (VPD), soil moisture deficit (for Mediterranean regions in particular) and phenology. The limiting effect of VPD on ozone uptake was especially apparent, since high VPDs resulting in stomatal closure tended to co-occur with high ozone concentrations. Although further work is needed to link the ozone uptake and deposition model components, and to validate the model with field measurements, the present results give a clear indication of the possible implications of adopting a flux-based approach for future policy evaluation.     

Persistent stimulation of CO2 assimilation and stomatal conductance by summer ozone fumigation in Norway spruce

CO(2) assimilation rate, stomatal conductance and chlorophyll content of current and previous years' needles of Norway spruce were measured in May 1988, 205 days after the cessation of ozone fumigation during the summer of 1987. Rates of assimilation were consistently higher for both needle year age classes for ozone fumigated trees in comparison to control trees, although only statistically significant for part of the day for current year's needles. A 26% and 48% stimulation, overall, in mean daily rates of assimilation for current and previous years' needles of ozone fumigated trees was observed. This was due to an enhanced apparent quantum yield and light saturated rate of assimilation of ozone fumigated trees. The temperature response regression of assimilation versus temperature was also greater, such that at any given temperature, assimilation was higher for ozone treated trees than control trees. Stomatal conductance was greater for ozone fumigated trees than the controls, but this was only marginally statistically significant. Moreover, there was a consistent increase in chlorophyll content in both year classes in ozone-treated trees. These results are discussed in relation to a possible long term effect of ozone fumigation upon the processes of conifer winter hardening and spring de-hardening.     

Photosynthetic and stomatal conductance responses of Norway spruce and beech to ozone, acid mist and frost--a conceptual model

Two-year-old beech and Norway spruce seedlings were exposed to a combination of ozone and acid mist treatments in open-top chambers in Scotland during the months of July through to September 1988. Replicate pairs of chambers received charcoal-filtered air (control), ozone-enriched air (140 nl ozone litre(-1)) or 140 nl ozone litre(-1) plus a synthetic acid mist (pH 2.5) composed of ammonium nitrate and sulphuric acid. Field measurements of assimilation and stomatal conductance were made during August. In addition, measurements of assimilation and conductance were made during September in the laboratory. Light response curves of assimilation and conductance were determined using a GENSTAT nonrectangular hyperbolic model. During February 1988/9 the Norway spruce were subject to a four day warming period at 12 degrees C and the light response of assimilation determined. The same plants were then subject to a 3-h night-time frost of -10 degrees C. The following day the time-course of the recovery of assimilation was determined. It was found that ozone fumigation did not influence the light response of assimilation of beech trees in the field, although stomatal conductance was reduced in the ozone-fumigated trees. The rate of light-saturated assimilation of Norway spruce was increased by ozone fumigation when measured in the field. Measurements of assimilation of Norway spruce made during the winter showed that prior to rewarming there was no difference in the rate of light-saturated assimilation for control and ozone-fumigated trees. However, the ozone plus acid mist-treated trees exhibited a significantly higher rate. The 4-day period of warming to 12 degrees C increased the rate of light-saturated assimilation in all treatments but only the ozone plus acid mist-treated trees showed a significant increase. Following a 3-h frost to -10 degrees C the control trees exhibited a reduction in the rate of light-saturated assimilation (Amax) to 80% of the pre-frost value. In comparison, following the frost, the ozone-fumigated trees showed an Amax of 74% of the pre-frost value. The ozone plus acid mist-treated trees showed an Amax of 64% of the pre-frost trees. The time taken for Amax to attain 50% of the pre-frost value increased from 30 min (control) to 85 min for ozone-fumigated trees to 190 min (ozone plus acid mist). These results are discussed in relation to the impact of mild, short-term frosts, which are known to occur with greater frequency than extreme, more catastrophic frost events. A simple conceptual framework is proposed to explain the variable results obtained in the literature with respect to the impact of ozone upon tree physiology.     

Comparison of different stomatal conductance algorithms for ozone flux modelling

A multiplicative and a semi-mechanistic, BWB-type [Ball, J.T., Woodrow, I.E., Berry, J.A., 1987. A model predicting stomatal conductance and its contribution to the control of photosynthesis under different environmental conditions. In: Biggens, J. (Ed.), Progress in Photosynthesis Research, vol. IV. Martinus Nijhoff, Dordrecht, pp. 221-224.] algorithm for calculating stomatal conductance (g(s)) at the leaf level have been parameterised for two crop and two tree species to test their use in regional scale ozone deposition modelling. The algorithms were tested against measured, site-specific data for durum wheat, grapevine, beech and birch of different European provenances. A direct comparison of both algorithms showed a similar performance in predicting hourly means and daily time-courses of g(s), whereas the multiplicative algorithm outperformed the BWB-type algorithm in modelling seasonal time-courses due to the inclusion of a phenology function. The re-parameterisation of the algorithms for local conditions in order to validate ozone deposition modelling on a European scale reveals the higher input requirements of the BWB-type algorithm as compared to the multiplicative algorithm because of the need of the former to model net photosynthesis (A(n)).     

A comparison of two stomatal conductance models for ozone flux modelling using data from two Brassica species

In this study we tested and compared a multiplicative stomatal model and a coupled semi-empirical stomatal-photosynthesis model in their ability to predict stomatal conductance to ozone (g_st) using leaf-level data from oilseed rape (Brassica napus L.) and broccoli (Brassica oleracea L. var. italica Plenck). For oilseed rape, the multiplicative model and the coupled model were able to explain 72% and 73% of the observed g_st variance, respectively. For broccoli, the models were able to explain 53% and 51% of the observed g_st variance, respectively. These results support the coupled semi-empirical stomatal-photosynthesis model as a valid alternative to the multiplicative stomatal model for O₃ flux modelling, in terms of predictive performance.     

Modelling stomatal ozone flux and deposition to grassland communities across Europe

Regional scale modelling of both ozone deposition and the risk of ozone impacts is poorly developed for grassland communities. This paper presents new predictions of stomatal ozone flux to grasslands at five different locations in Europe, using a mechanistic model of canopy development for productive grasslands to generate time series of leaf area index and soil water potential as inputs to the stomatal component of the DO(3)SE ozone deposition model. The parameterisation of both models was based on Lolium perenne, a dominant species of productive pasture in Europe. The modelled seasonal time course of stomatal ozone flux to both the whole canopy and to upper leaves showed large differences between climatic zones, which depended on the timing of the start of the growing season, the effect of soil water potential, and the frequency of hay cuts. Values of modelled accumulated flux indices and the AOT40 index showed a five-fold difference between locations, but the locations with the highest flux differed depending on the index used; the period contributing to the accumulation of AOT40 did not always coincide with the modelled period of active ozone canopy uptake. Use of a fixed seasonal profile of leaf area index in the flux model produced very different estimates of annual accumulated total canopy and leaf ozone flux when compared with the flux model linked to a simulation of canopy growth. Regional scale model estimates of both the risks of ozone impacts and of total ozone deposition will be inaccurate unless the effects of climate and management in modifying grass canopy growth are incorporated.     

Modelling annual pasture dynamics: Application to stomatal ozone deposition

Modelling ozone (O₃) deposition for impact risk assessment is still poorly developed for herbaceous vegetation, particularly for Mediterranean annual pastures. High inter-annual climatic variability in the Mediterranean area makes it difficult to develop models characterizing gas exchange behaviour and air pollutant absorption suitable for risk assessment. This paper presents a new model to estimate stomatal conductance (g_s) of Trifolium subterraneum, a characteristic species of dehesa pastures. The MEDPAS (MEDiterranean PAStures) model couples 3 modules estimating soil water content (SWC), vegetation growth and g_s. The g_s module is a reparameterized version of the stomatal component of the EMEP DO₃SE O₃ deposition model. The MEDPAS model was applied to two contrasting years representing typical dry and humid springs respectively and with different O₃ exposures. The MEDPAS model reproduced realistically the g_s seasonal and inter-annual variations observed in the field. SWC was identified as the major driver of differences across years. Despite the higher O₃ exposure in the dry year, meteorological conditions favoured 2.1 times higher g_s and 56 day longer growing season in the humid year compared to the dry year. This resulted in higher ozone fluxes absorbed by T. subterraneum in the humid year. High inter-family variability was found in gas exchange rates, therefore limiting the relevance of single species O₃ deposition flux modelling for dehesa pastures. Stomatal conductance dynamics at the canopy level need to be considered for more accurate O₃ flux modelling for present and future climate scenarios in the Mediterranean area.     

The impact of constituent ions of acid mist on assimilation and stomatal conductance of Norway spruce prior and post mid-winter freezing

Norway spruce seedlings were sprayed twice weekly with one of a range of artificial mists at either pH 2.5, 3.0 or 5.6, for three months. The mists consisted of either (NH4)2SO4 (pH 5.6), NH4NO3 (pH 5.6), water (pH 5.6), HNO3 (pH 2.5), H2SO4 (pH 2.5). In late December 1988 and early January 1989 the light response of assimilation and stomatal conductance were assessed in the laboratory following a 4-day equilibration period at 12 degrees C. The intact trees were then subjected to a mild (-10 degrees C), brief (3 h) frost in the dark and the recovery of light saturated assimilation (Amax) was followed during the subsequent light period. The same trees were then subjected to a second 3 h (-18 degrees C) frost. The recovery of Amax during the next day was followed. All ion-containing mists stimulated Amax and apparent quantum yield relative to control trees, irrespective of pH. The mists containing SO4 made stomatal conductance unresponsive to light flux density and caused the stomata to lock open. Frosts of -10 degrees C and -18 degrees C did not inhibit the Amax of control trees for longer than 200 min into the light period. In contrast, the ion-containing mists exerted a significant inhibitory effect upon the recovery of Amax. Nitric acid inhibited Amax to 35% of the pre-frost value, whilst the remaining treatments inhibited Amax between 15% and 40% of the pre-frost value. It is concluded that SO4 causes increased mid-winter frost sensitivity and NO3 ameliortes this effect. The results are discussed in relation to forest decline.     

Limited compensation of ozone stress by potassium in Norway spruce

The effects of potassium fertilization and ozone stress were investigated in a clone of Picea abies (L.) Karst, by studies of the uptake of CO(2) by the crowns, the element content, on leaching of the youngest needles, and the longevity of the needles. All plants were exposed to 0.075 microl litre(-1) SO(2) from January to April 1986. The average ozone concentrations applied during the subsequent growing season (May-December) were 0, 0.027, 0.050 and 0.100 microl litre(-1). Half of the trees received liquid fertilizer applications from April to July 1986. CO(2) uptake by the crowns was significantly reduced in non-fertilized plants at ozone doses of 100-200 microl litre(-1) h, whereas similar reductions were recorded in fertilized plants only above an ozone dose of 300 microl litre(-1) h. Independent of the fertilization, however, the concentrations of calcium, magnesium and nitrogen in the needles increased in parallel with the ozone dose, whilst potassium, phosphorus and sulphur showed little response to ozone. In both nutrient regimes, the diffusive loss of elements from chloroform-washed needles was similar and tended to be reduced at the highest ozone concentration, when relating the leachate to the corresponding element content in the needles. Needles formed in the highest ozone treatment were significantly shed during the succeeding year, regardless of the nutrient supply. It appears that increased potassium supply has little compensating effect on ozone stress in spruce.     

Ozone flux over a Norway spruce forest and correlation with net ecosystem production

Daily ozone deposition flux to a Norway spruce forest in Czech Republic was measured using the gradient method in July and August 2008. Results were in good agreement with a deposition flux model. The mean daily stomatal uptake of ozone was around 47% of total deposition. Average deposition velocity was 0.39 cm s⁻¹ and 0.36 cm s⁻¹ by the gradient method and the deposition model, respectively. Measured and modelled non-stomatal uptake was around 0.2 cm s⁻¹. In addition, net ecosystem production (NEP) was measured by using Eddy Covariance and correlations with O₃ concentrations at 15 m a.g.l., total deposition and stomatal uptake were tested. Total deposition and stomatal uptake of ozone significantly decreased NEP, especially by high intensities of solar radiation.     

Simulation of stomatal conductance for Aleppo pine to estimate its ozone uptake

The data from a previous experiment carried out in open-top chambers to assess the effects of ozone (O3) exposure on growth and physiology of Aleppo pine (Pinus halepensis Mill.) were re-assessed to test the performance of the EMEP O3 stomatal conductance model used to estimate tree O3 uptake at a European scale. Aleppo pine seedlings were exposed during three consecutive years to three different O3 treatments: charcoal filtered air, non-filtered air and non-filtered air supplemented with 40 nl l(-1). The results of the model using the default parameterisation already published for Mediterranean conifers showed a poor performance when compared to measured data. Therefore, modifications of g(max), f(min), and new f(VPD), f(temp) and f(phen) functions were developed according to the observed data. This re-parameterisation resulted in a significant improvement of the performance of the model when compared to its original version.     

Leaf conductance response of phaseolus vulgaris to ozone flux density

The effect of ozone flux density on leaf conductance to ozone in Phaseolus vulgaris was examined. The change in conductance was measured within the first two hours of fumigation for mature, fruiting 6-week-old plants of an ozone sensitive cultivar (Seafarer); for young, 14-day-old plants of the same cultivar; and for an ozone resistant cultivar (Gold Crop). Young Seafarer plants showed no change in conductance to ozone over a wide range of ozone flux densities. Gold Crop showed a decrease in conductance of −3.1 % /(mgO₃ m⁻² h⁻¹) whereas mature Seafarer plants exhibited a stronger decrease of −7.7% /(mgO₃ m⁻² h⁻¹). Diffusion porometer measurements taken on fruiting Seafarer plants in the field illustrated that a decrease in leaf diffusive conductance to water is related to visual ozone injury.     

Persistent effects of ozone on needle water loss and wettability in Norway spruce

Four-year-old, seed-grown trees of Norway spruce (Picea abies (L.) Karst.) were exposed in open-top chambers to charcoal-filtered air (8 h daily mean 54 microg O(3) m(-3)) over three consecutive summers (1986-1988). In mid-May 1988, before the third season of fumigation and more than 7 months after exposure to ozone the previous summer had terminated, daily rates of transpiration from intact shoots and water loss from excised needles were measured together with the amount of wax on the needle surface. In mid-July, 92 days after the beginning of the third year of exposure, the wettability of needles was assessed by measuring the contact angle of water droplets on the surface of needles. Exposure to 156 microg O(3) m(-3) resulted in a 16% increase in daily transpiration in current year's needles and a 28% increase in 1-year old needles. These effects were associated with slower stomatal closure in response to increasing water deficit in the needles previously exposed to 156 microg m(-3) ozone. The long-lasting nature of such ozone-induced effects could predispose trees to drought and winter desiccation. No significant effects of ozone were found on the amount of wax covering the needle surface, but a marked increase in the wettability of needles exposed to ozone was observed. The far reaching physiological consequences of these effects in the field and the possibility that similar disturbances may contribute to the decline of high-altitude forests of Norway spruce in Europe are discussed.     

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.     

Tree and stand growth of mature Norway spruce and European beech under long-term ozone fumigation

In a 50- to 70-year-old mixed stand of Norway spruce (Picea abies (L.) Karst.) and European beech (Fagus sylvatica L.) in Germany, tree cohorts have been exposed to double ambient ozone (2×O₃) from 2000 through 2007 and can be compared with trees in the same stand under the ambient ozone regime (1×O₃). Annual diameter growth, allocation pattern, stem form, and stem volume were quantified at the individual tree and stand level. Ozone fumigation induced a shift in the resource allocation into height growth at the expense of diameter growth. This change in allometry leads to rather cone-shaped stem forms and reduced stem stability in the case of spruce, and even neiloidal stem shapes in the case of beech. Neglect of such ozone-induced changes in stem shape may lead to a flawed estimation of volume growth. On the stand level, 2×O₃ caused, on average, a decrease of 10.2 m³ ha⁻¹ yr⁻¹ in European beech.     

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%.     

Deposition of ozone to a mature spruce forest: measurements and comparison to models

Removal of ozone at terrestrial surfaces provides a major sink for tropospheric ozone and, therefore, a constraint on the peak concentrations achieved during photochemical episodes. This study reports results from 5 years of almost continuous measurements of vertical profiles of ozone and related meteorological variables over a mature spruce forest in Bavaria. Deposition velocities calculated from flux/gradient and eddy correlation flux measurements have been compared with estimates based on a resistance model and yield satisfactory agreement during fine weather conditions. The results also suggest that biogenic emissions of reactive hydrocarbons from the forest influence the vertical profile of ozone.     

Implications of climate change for the stomatal flux of ozone: a case study for winter wheat

Climate change factors such as elevated CO2 concentrations, warming and changes in precipitation affect the stomatal flux of ozone (O3) into leaves directly or indirectly by altering the stomatal conductance, atmospheric O3 concentrations, frequency and extent of pollution episodes and length of the growing season. Results of a case study for winter wheat indicate that in a future climate the exceedance of the flux-based critical level of O3 might be reduced across Europe, even when taking into account an increase in tropospheric background O3 concentration. In contrast, the exceedance of the concentration-based critical level of O3 will increase with the projected increase in tropospheric background O3 concentration. The influence of climate change should be considered when predicting the future effects of O3 on vegetation. There is a clear need for multi-factorial, open-air experiments to provide more realistic information for O3 flux-effect modelling in a future climate.     

The use of light and electron microscopy to assess the impact of ozone on Norway spruce needles

Light (LM) and transmission electron (TEM) microscopy were used to study previously specified ozone symptoms in the foliage of Norway spruce. The three youngest green needle generations from twenty mature trees in two stands on sites of different soil fertility at Asa, southern Sweden, were sampled in 1999. The critical dose of ozone, expressed as AOT40, was 6,362 ppb.h. LM showed ozone-specific symptoms: decreased chloroplast size with electron dense stroma advancing gradually from the outer to the inner cell layers, being most severe in the needle side facing the sky. The symptoms were expressed as ozone syndrome indices at the needle generation, tree and stand levels. The index had higher values at the low fertility site. TEM was used to confirm the LM results. The study shows that LM can be used for diagnosis of the impact of ozone on conifers in the field.