Modelling stomatal ozone ux across Europe

L.D. Emberson
a,b,
*, M.R. Ashmore
b
, H.M. Cambridge
a
, D. Simpson
c
, J.-P. Tuovinen
d
a
Biology Department, Stockholm Environment Institute at York, University of York, Box 373, York YO10 5YW, UK
b
Department of Environmental Science, University of Bradford, Bradford, West Yorkshire, BD7 1DP, UK
c
EMEP MSC-W, Norwegian Meteorological Institute, Oslo, Norway
d
Finnish Meteorological Institute, Sahaajankatu 20 E, FIN-00810, Helsinki, Finland
Received 30 June 1999; accepted 5 January 2000
``Capsule'': The stomatal ux of ozone for dierent types of vegetation is modelled on a European scale; dierences in
the spatial distribution between stomatal ux and the exposure index AOT40 indicate important implications for risk
analysis.
Abstract
A model has been developed to estimate stomatal ozone ux 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 ux 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 eect of phenology, irradiance, temperature, vapour
pressure decit and soil moisture decit on stomatal conductance. These species-specic 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 uxes are presented as mean monthly ux maps and
compared with maps describing equivalent values of AOT40 (accumulated exposure over threshold of 40 ppb or nl l
1
), high-
lighting the spatial dierences between these two indices. In many cases high ozone uxes were modelled in association with only
moderate AOT40 values. The factors most important in limiting ozone uptake under the model assumptions were vapour pressure
decit (VPD), soil moisture decit (for Mediterranean regions in particular) and phenology. The limiting eect 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 eld
measurements, the present results give a clear indication of the possible implications of adopting a ux-based approach for future
policy evaluation. #2000 Elsevier Science Ltd. All rights reserved.
Keywords: Ozone; Ozone ux; Stomatal conductance; Critical levels; Micrometeorology; Risk analysis
1. Introduction
Current European levels of tropospheric ozone have
been shown to cause damage to forest trees, agricultural
crops and semi-natural vegetation (Ka renlampi and
Ska rby, 1996). In order to address this, the United
Nations Economic Commission for Europe (UN-ECE)
has adopted an eects-based approach, using the critical
loads/levels concept. Such critical levels for ozone will
play an important role in the multi-pollutant/multi-
eect protocols which are being developed to aid policy
formulation to control ozone precursor emissions and
reduce European ozone concentrations to acceptable
levels (Bull, 1991). The existing eects-based research
for ozone has resulted in the establishment of critical
levels to a so-called Level I standard (Ka renlampi and
Ska rby, 1996). These Level I values relate ozone injury
to ozone exposure using a cumulative exposure over a
threshold concentration of 40 ppb (nl l
1
) for daylight
hours (AOT40), with the necessary exposureresponse
relationships being derived from open-top chamber
experiments. These experiments demonstrated linear
relationships between cumulative seasonal ozone expo-
sure, expressed as AOT40, and a variety of plant
responses (Fuhrer et al., 1997). From these relationships
it has been possible to derive a critical level based on a
0269-7491/00/$ - see front matter # 2000 Elsevier Science Ltd. All rights reserved.
PI I : S0269- 7491( 00) 00043- 9
Environmental Pollution 109 (2000) 403413
www.elsevier.com/locate/envpol
* Corresponding author. Tel.: +44-1904-432896; fax: +44-1904-
432898.
E-mail address: l.emberson@york.ac.uk (L.D. Emberson).
given level of plant response (e.g. a 5% yield reduction).
However, extrapolating from these chamber experi-
ments to ambient conditions is dicult for two reasons.
Firstly, the dierences in microclimate between the
chamber-grown plants and those growing outside may
lead to dierences in plant response to the same ozone
exposure (e.g. Sanders et al., 1991; Pleijel et al., 1994),
because of dierences in aerodynamic and boundary
layer conductances, and dierences in plant sensitivity.
Secondly, the maintenance of chamber plants, e.g.
through watering or the application of systemic fungi-
cides, may alter plant sensitivity to air pollutants. These
factors create uncertainty in the value of the Level I
critical levels that have been proposed to protect vege-
tation.
It is also important to note that the exposure
response relationships from which the Level I critical
level values have been derived are based on applying a
range of dierent ozone exposures to replicate chambers
in the same location and under the same climatic con-
ditions. It has been emphasised by a number of authors
(e.g. Fuhrer et al., 1997; Gru nhage and Haenel, 1997)
that these chamber-based relationships cannot be used
to assess the relative size of ozone impacts at dierent
locations across Europe. In particular, it has long been
recognised that plant response is more closely related to
the internal ozone dose, or the instantaneous ux of
ozone through the stomata, than the ambient ozone
exposure (e.g. Amiro et al., 1984; Fuhrer et al., 1992). It
is important to note that high concentrations of ozone
are often associated with factors leading to reduced
ozone ux, such as high vapour pressure decits (VPDs)
(Gru nhage and Ja ger, 1994; Gru nhage et al., 1997).
Furthermore, the highest ozone concentrations in areas
such as southern Europe occur during seasons when
non-irrigated vegetation experiences high soil moisture
decits (SMDs), resulting in reduced stomatal con-
ductance and hence low ozone ux.
As such, the Level I critical level values, which are
being used in developing European ozone control stra-
tegies both by UN-ECE and the European Union (EU),
are intended to protect the most sensitive vegetation
types under the most sensitive conditions. Exceedance
of these critical levels only provides an indication that
some risk exists of damage to vegetation from ozone;
the degree of exceedance cannot be used to provide a
measure of the relative risk of damage to vegetation in
dierent areas of Europe.
It is thus clear that an assessment based on ozone ux
to receptor sites within the leaf, rather than ozone
exposure, could provide an improved estimate of the
relative degree of risk of ozone damage to vegetation
across Europe, and hence allow more cost-eective
control strategies to be identied. Existing models of
ozone ux (e.g. Baldocchi et al., 1987; Ko rner et al.,
1995; Gru nhage and Haenel, 1997) require detailed
micrometeorological information, or are applicable to
only a limited number of species, and cannot be readily
applied at a European scale. Alternatively, models
designed to model ozone deposition on a regional scale
(e.g. Wesely, 1989; Erisman et al., 1994) have a limited
description of the stomatal responses, which are not
species-specic and which do not consider the eects of
VPD or SMD.
Our objectives in this study were to develop a model-
ling approach, which could be applied to estimate and
map stomatal ozone ux to major vegetation types
across Europe. The methodology we report in this paper
enables the calculation of ozone ux to physiologically
active leaves of the upper canopy as a function of ozone
exposure, meteorology, stomatal function and soil data.
This allows the importance of dierent model para-
meters in determining ozone uptake to be estimated,
and enables comparisons of the temporal and spatial
variation in modelled ozone uxes with those of the
AOT40 index. Application of the model over such a
large spatial scale gives an indication of the temporal
and spatial variability of absorbed ozone dose, taking
into account the systemic spatial variation in climate
across Europe. It should be noted that the development
of this model has largely concentrated on the stomatal
component of ozone uptake since this is recognised as
the predominant factor limiting ozone ux into the
leaves. However, the atmospheric and boundary layer
components are included in a simplied manner to give
an indication of the relative eect of these resistances on
ozone uptake. Thus, although the present calculations
can be expected to improve in future, the current
approach allows us for the rst time to present regional-
scale comparisons of modelled stomatal ux for two
major species with the ozone indices currently used to
determine ozone risk. Such comparisons give a clear
indication of the possible implications of adopting a
ux-based approach for future policy evaluation.
2. Methodology
The modelling approach for stomatal resistance (r
s
) is
based on the multiplicative stomatal conductance model
(where conductance is the reciprocal of resistance)
described by Jarvis (1976) and modied by Ko rner et al.
(1995). This model has been further developed to enable
the evaluation of species-specic stomatal conductance
(g
s
) on a European scale. The model has been para-
meterised for a number of dierent European species,
including 10 tree species, 7 agricultural crops and 1 type
of semi-natural vegetation. This has been achieved
through collecting and pooling information available
from the scientic literature. To ensure comparability of
the collected g
s
data rigorous criteria were set regarding
the procurement and recording of the data before they
404 L.D. Emberson et al. / Environmental Pollution 109 (2000) 403413
were accepted for use in establishing the g
s
dataset
(Emberson, 1997).
Stomatal conductance was calculated as a function of
species type, phenology and environmental conditions
[photon ux density (PFD), temperature, VPD, SMD]
since these are the factors considered most important in
determining stomatal aperture. This basic model has
been developed to calculate species-specic g
s
according
to phenology and prevailing environmental conditions
with g
s
responding to these factors as described in
Eq. (1).
g
s
= g
max
g
pot
max{g
min
; (g
light
g
temp
g
VPD
g
SWP
)]; (1)
where g
max
is the species-specic maximum stomatal
conductance to ozone (mmol m
2
s
1
) expressed on a
total leaf surface area. The parameters g
pot
, g
light
, g
temp
,
g
VPD
and g
SMD
are all expressed in relative terms (i.e.
they take a value between 0 and 1) as a proportion of
g
max
where:
g
pot
represents the modication to g
max
due to phe-
nological changes;
g
light
represents the modication of g
max
by PFD
(mmol m
2
s
1
);
g
temp
represents the modication of g
max
by leaf tem-
perature (

C);
g
VPD
represents the modication of g
max
by VPD
(kPa);
g
SWP
represents the modication of g
max
by SMD
(MPa); and
g
min
represents the minimum stomatal conductance
that occurs during the daylight period. As such, g
temp
,
g
VPD
and g
SMD
are set equal to g
min
if the values
calculated in Eqs. (5), (6) and (7) fall below g
min
.
The model development described here is unique to
this study because of the parameterisation of the g
s
relationships for a large number of important European
species. In addition, the model relates g
s
to soil moisture
status rather than leaf water status, and is combined
with a model capable of evaluating species-specic
SMD. These additions to the model make it possible to
calculate g
s
for a number of dierent species across the
whole of Europe. The pan-European nature of the study
means that the development and application of the
methodology is to some extent limited by the avail-
ability of input data. For this initial application of the
deposition model, the Norwegian Meteorological Insti-
tute (DNMI) provided European meteorological data
for the year 1994 at a spatial scale of 150150 km and a
temporal scale of 6 h. These meteorological data were
manipulated for dierent land-cover types within a grid
to derive the necessary input environmental parameters
representing the species-specic microclimatic con-
ditions as discussed below. In the case of SMD, the
evaluation of soil moisture is land-cover specic.
The response of stomatal conductance to phenology is
accounted for by the species-specic g
pot
functions
which modify g
max
to give a potential maximum value
of g
s
according to the day within the growing season for
each species, assuming linear increases and decreases at
the start and end of the growing season, with a period
when g
pot
is 1 in the middle of the growing season g
pot
is
thus modelled as follows:
g
pot
= (1 g
pot a
) ((yd SGS)=g
pot b
) g
pot a
when SGS4yd4(SGS g
pot b
);
g
pot
= 1 (2)
when (SGS g
pot b
)4yd4(EGS g
pot c
);
g
pot
= (1 g
pot a
) ((EGS yd)=g
pot c
) g
pot a
when EGS5yd5(EGS g
pot c
);
where g
pot_a
, g
pot_b
and g
pot_c
are species-specic para-
meters representing, in turn, the minimum g
pot
, the
number of days for g
pot
to reach its maximum and the
number of days during the decline of g
pot
for the mini-
mum to again be reached. Yd is the year day and SGS
and EGS represent the species-specic start and end of
the growing season, respectively.
The beech growing season was estimated using a
thermal time model to calculate the eective temperature
sum (ETS) and predict leaf emergence (Kramer, 1994):
ETS (n) =
X
n
i=1
max(0; T
i
T
b
) (3)
where n is the year day, T
i
is the mean daily temperature
and T
b
a pre-dened threshold value (4.5

C). The
critical ETS for budburst is 206.4 degree-days and the
year day at which this ETS is reached represents SGS
(Kramer, 1994). The date of beech leaf fall across
Europe was assumed constant at year day 306 and
represents EGS (Kramer, 1995).
The wheat-growing season was calculated using a
latitude model established by collecting wheat leaf
emergence data from a number of dierent studies
across Europe (Peterson, 1965; Lo vblad et al, 1996) and
relating this to latitude across a range of 3070

. A
standard growing season length of 3 months is assumed.
This is consistent with the growing season over which
L.D. Emberson et al. / Environmental Pollution 109 (2000) 403413 405
the AOT40 for spring wheat is currently calculated for
Level I ozone critical levels.
The response of stomatal conductance to PFD in
units of mmol m
2
s
1
is estimated using a rectangular
hyperbola:-
g
light
= 1 exp
light aPFD
; (4)
where light_a is a species-specic parameter dictating the
shape of the hyperbolic relationship. PFD is calculated
using a standard European Monitoring and Evaluation
Programme (EMEP) procedure which has been used
previously in dry deposition models, as described by
Jakobsen et al. (1996). The method was devised to esti-
mate PDFas a function of cloud cover, latitude, longitude
and time, and uses established solar geometry techniques.
The stomatal response to leaf temperature (T), in
degrees Celcius, is described by a parabolic function:
g
temp
= 1 (T T opt)
2
=(T opt T min)
2
(5)
where T
min
is the minimum temperature at which sto-
matal opening occurs and T
opt
is the optimumtemperature.
DNMI meteorological data provide air temperature
data at a height of 2-m above the ground surface. The
temperature of a leaf in the upper canopy is calculated
from the 2-m air temperature data as a function of sen-
sible heat ux density and leaf dimension (Jones, 1992).
This calculation is only used when global radiation levels
are higher than 50 W m
2
; under other conditions, the leaf
temperature is assumed equal to the air temperature.
The stomatal response to VPD between the thresholds
for minimum and full stomatal opening (represented by
VPD_min and VPD_max, respectively, in kPa) is
described by the relationship:
g
VPD
= min{1; ((1 g
min
) (VPD min VPD)=
(VPD min VPD max)) g
min
]:
(6)
For this initial application of the deposition model,
VPD was calculated assuming that specic humidity is
constant across the atmospheric boundary layer, allow-
ing the 2 m height humidity to be calculated according
to Monteith and Unsworth (1990). This value is con-
verted to VPD using a standard temperature function to
calculate relative leaf to air VPD (Jones, 1992) based on
the model leaf temperatures.
The stomatal response to SMD between thresholds
for minimum stomatal opening and full stomatal open-
ing (represented by SWP_min and SWP_max, respec-
tively) is described by the relationship:
g
SWP
= min{1; ((1 g
min
) (SWP min
SWP=(SWP min SWP max)) g
min
]: (7)
The g
SWP
values of SWP_min and SWP_max are
collected from literature describing stomatal responses
to either increasing soil or predawn leaf water potential.
The conversion to units of volumetric water content was
achieved using soil water release curves. Three curves
were established for each of the three soil textures
(coarse, medium, ne) mapped across Europe according
to soil texture data (FAO-UNESCO, 1981). Functions
for each soil texture were established using the relevant
physical characteristics of the dierent soil textures
described by the Food and Agriculture Organisation
(FAO), in combination with functions dened by
Campbell (1985) and Milthorpe and Moorby (1974).
The available soil water (ASW) parameter for each soil
type was calculated according to species type, using the
g
SWP
functions collected from the literature (Table 1)
where permanent wilting point is assumed to occur at
SWP_min. Field capacity is assumed to be constant
within a soil type and equivalent to 0.01 for coarse
and 0.005 MPa for medium- and ne-textured soils,
respectively. The species-specic g
SWP
function, in units
of MPa, was converted to an equivalent function in
units of SMD using the soil water release curves and
species- and soil texture- specic rooting depths
(Thornthwaite and Mather, 1957; Jackson et al., 1996).
This enables the eect of SMD on g
s
to be calculated.
The calculation of SMD used DNMI meteorological
data. Surface temperature and precipitation data were
used to calculate potential evapotranspiration. Actual
evapotranspiration was evaluated according to a proce-
dure established by Thornthwaite and Mather (1957)
and modied to incorporate the species-specic ASW.
Thornthwaite and Mather's (1957) method is internally
consistent in that the calculation of SMD is made with
reference to an original soil water status. SMD was cal-
culated from the start of 1994 using the DNMI input
data. However, SMD is a cumulative variable, and
therefore a starting SMD value is required to describe
the soil water status at the beginning of the 1994 period.
It would not be correct to assume that across the whole
of Europe SMD is equal to zero since some Mediterra-
nean areas are known to rarely, if ever, reach eld
capacity by the beginning of January. To overcome this
problem, SMD was modelled using 30-year average
data (Cramer, personal communication) using the same
method described above, to calculate an original SMD
which is assumed to represent the soil water status for
the beginning of 1994.
3. Model application
The methodology described above has been applied
on a pan-European scale. Model runs have been per-
formed for the duration of the species-specic growing
season during 1994. The results are described here to
406 L.D. Emberson et al. / Environmental Pollution 109 (2000) 403413
show the potential applications of the model and also to
give an indication of the spatial and temporal variation
between this ux modelling approach and the existing
Level I exposure approach. These runs are performed
for only two species, beech (Fagus sylvatica) and spring
wheat (Triticum aestivum) for which the g
s
para-
meterisations are given in Table 1. However, it is
stressed that the capability exists to perform this
modelling for any of the forest tree, agricultural crops
or semi-natural vegetation types for which g
s
parameterisation have been collected. Beech and wheat
were selected for this preliminary investigation due
to their wide distribution across Europe, and since they
are the species upon which the current critical Level I
values are based for forest trees and agricultural crops
respectively.
The model runs performed in this study use meteor-
ology and ozone data from the Lagrangian EMEP
photo-oxidant model (Simpson, 1993, 1995). This model
predicts mean ozone concentrations over the atmo-
spheric boundary layer, typically ca. 1 km deep. These
values have been used to estimate near-surface con-
centrations of ozone using similarity theory (Simpson,
1993), applied to each land-cover class within the grid,
assuming a surface layer depth of 50 m.
Since ozone uptake is not only dependent upon the
species-specic stomatal conductance, but also upon the
atmospheric and boundary layer conductance to ozone,
these factors need to be incorporated in an assessment
of stomatal ozone ux. In addition, the concentration at
the leaf surface is also dependent upon the ozone
deposition to the canopy as a whole, since this deter-
mines the ozone concentration available for stomatal
uptake by the plant. These non-stomatal pathways have
been neglected in the present study hence these initial
calculations probably overestimate the near-surface
ozone and hence stomatal uptake. However, this uncer-
tainty is minimised to some extent in this modelling
study, since ozone uptake is calculated for a leaf in the
upper canopy. The accuracy of the calculation of ozone
ux could be improved and used to describe canopy
uptake if Leaf Area Index data were combined with the
rate of ozone deposition to the plant cuticle, and
through the canopy to the underlying soil surface.
However, since the parameterisation for such a com-
plex deposition/stomatal ux model is not at present
available, we have combined the atmospheric, boundary
layer and stomatal components in a simplied manner.
The methods used allow us to gain an indication of
the relative importance of the factors determining
ozone uptake and how these might vary diurnally
throughout the growing season with variable ozone
concentrations.
The simplied model is as follows:
GO
3spp
=
1
Ra
spp
rb
spp
rs
spp
; (8)
Table 1
Parameters used to calculate leaf stomatal conductance for wheat and beech indicating key literature sources
Parameter Beech Beech references Wheat Wheat references
g
max
(mmol O
3
m
2
s
1
)
a
66 Pearson and Manseld (1993); Mikkelsen
(1995)
154 Araus and Tapia (1987); Lehnherr et al.
(1988); Araus et al. (1989); Grandjean Grimm
and Fuhrer (1992a); Gru ters et al. (1995)
g
min
0.13 Tognetti et al. (1995) 0.1 Ko rner (1994)
g
pot
_a 0.3 Taylor and Dobson (1989);
Pearson and Manseld (1993)
0.1 Araus and Tapia (1987); Grandjean Grimm
and Fuhrer (1992b); Gru ters et al. (1995)
g
pot
_b (days) 50 0
g
pot
_c (days) 50 45
Light_a 0.006 Eamus and Murray (1991) 0.009 Machado and Lago (1994); Gru ters et al.
(1995); Weber and Rennenberg (1996)
T_max (

C) 34 Osonubi and Davies (1980); Matyssek

(personal communication)
40 Gru ters et al. (1995)
T_min (

C) 13 13
T_opt (

C) 24 26
VPD_max (kPa) 1.1 Kerstiens (1995) 0.9 Tuebner (1985); Weber and Rennenberg
(1996)
VPD_min (kPa) 3.1 2.8
SWP_max (MPa) 1.0 Tognetti et al. (1995) 0.3 Morgan (1984); Gollan et al. (1986);
Emberson (1997)
SWP_min (MPa) 1.9 1.1
a
Based on total leaf area.
L.D. Emberson et al. / Environmental Pollution 109 (2000) 403413 407
where Ra
spp
is the bulk atmospheric resistance to the
canopy as a whole and rb
spp
and rs
spp
are the species-spe-
cic boundary layer and stomatal resistances to an indi-
vidual leaf of the upper canopy respectively. This value is
then used in the calculation of ozone uptake as described
in Eq. (9), which assumes that there are reactive sinks
within the leaf that reduce the internal ozone concentra-
tion to zero (Laisk et al., 1989; Wang et al., 1995).
O
3
flux (nmol m
2
s
1
) = GO
3
(mol m
2
s
1
)
O
3
concentration at 50 m height(nmol mol
1
) (9)
The mean aerodynamic resistance for each grid
square is calculated as the resistance to mass transfer
exerted by turbulence between the top of the surface
layer (assumed 50 m in the EMEP model) and the
vegetation surface. The calculation of Ra follows that
described by Jakobsen et al. (1996) which calculates
species-specic bulk atmospheric resistance (Ra
spp
) as a
function of canopy height and roughness length. The
boundary layer resistance (rb
spp
) represents the limita-
tion to ozone transfer resulting from quasi-laminar
boundary layer found adjacent to the leaf surface.
Across this air layer, ozone transport occurs pre-
dominantly via molecular diusion, as opposed to the
mass transport associated with the aerodynamic com-
ponent. rb
spp
is a function of wind speed and surface
properties and is calculated as shown in Eq. (10), for a
single leaf of the upper canopy (e.g. Hosker, 1986).
rb
spp
=
d=(DO
3
ShO
3
)
2
; (10)
where ShO
3
is the Sherwood number for ozone for a at
leaf calculated as a function of near surface windspeed,
leaf dimensions and empirically derived constants
(Monteith and Unsworth, 1990). DO
3
is the molecular
diusivity of ozone in air (0.15 cm
2
s
1
) and d is the
mean leaf length in the direction of the wind.
To enable the modelling results to be presented as
ozone ux maps the locations of both species need to be
identied across Europe. Species location is dened
according to land-cover data, which were compiled and
manipulated at the Stockholm Environment Institute
(SEI) to give percentage land-cover values of each species
within each of the 150150-km grid squares. Beech
land-cover was obtained from the Forest Map of
Europe (ESA, 1992) and the FAO Land Use Map
of Europe (FAO-Cartographia, 1980). The distribution
of wheat was obtained using the FAO Land Use Map of
Europe (FAO-Cartographia, 1980) to delimit the agri-
cultural land area of Europe. Agricultural type within
this land area was obtained from Kostrowski (1984)
Types of Agriculture Map in combination with statis-
tical information describing wheat yield data (Eurostat,
1994). These statistical data enabled identication of
those land areas where wheat was the dominant crop
type. The modelling for wheat and beech was performed
within each grid square for which species occurrence
was predicted.
The input data used for this preliminary modelling
analysis used meteorological and ozone concentration
data at a 6 hourly temporal resolution. Therefore, the
results relate to 0, 6, 12 and 18 h over the diurnal period
throughout the growing season for both species. This
information is presented in a number of dierent ways
depending upon the aspect of ozone uptake under
investigation.
4. Results and discussion
The modelling procedure was used to generate maps
of monthly mean ozone uxes for both beech and wheat
across Europe. These ux maps were compared with
equivalent AOT40 maps in order to compare the spatial
variability between the ux and exposure-based risk
assessment approaches. Figs. 13 show the AOT40 and
ux maps for both wheat and beech, for June during
1994.
One of the main ndings from the comparisons of the
ux and exposure maps were that the areas experiencing
the highest ozone exposures (i.e. where the AOT40
values were highest) were frequently not the same as
those regions calculated as having the highest ozone
uxes. Fig. 1 shows the highest ozone exposures
(AOT40 values) to occur in the Mediterranean region
and central Europe. In contrast, it is apparent that for
wheat (Fig. 2), the highest ozone uxes occur in
southern Scandinavia and northern Europe with uxes
ranging from 1.5 to 2 nmol m
2
s
1
. The wheat ux
map also shows distinct bands of ozone ux of similar
classes across Europe, with the lowest classes occurring
in the more southerly latitudes. This is a result of the
growing season occurring earlier at the lower latitudes
and hence the earlier onset of a decrease in g
pot
limiting
stomatal conductance and hence ozone uptake. For
beech (Fig. 3), the dierences in spatial pattern between
exposure and ux are not so great, although the more
moderate AOT40 values found on the outskirts of
mainland Europe coincide with some of the highest
uxes recorded across Europe with values of 1 to 1.25
nmol m
2
s
1
.
To investigate the relationships between the dierent
components associated with ozone uptake, and to com-
pare regional ozone uxes and exposures over the entire
growing season for both beech and wheat, four EMEP
grid squares were selected for further analysis of the
modelled data. These grid squares were located in
southern Sweden, UK, Czech Republic and southern
Spain, and were selected to represent climatically
408 L.D. Emberson et al. / Environmental Pollution 109 (2000) 403413
diverse regions of Europe, each experiencing dierent
ozone exposure regimes.
Table 2 compares the AOT40 and ux values in these
squares for wheat and beech during 1994. Comparisons
are made between total cumulative uptake values
(CUO
3
in mmol m
2
), cumulative uptake values above
an arbitrary threshold of 1.5 nmol O
3
m
2
s
1
(TCUO
3
in mmol m
2
) and AOT40 (in ppb.h), calculated over
the growing season of the crop. The arbitrarily selected
ux threshold of 1.5 nmol m
2
s
1
(expressed on a total
leaf area basis) is equivalent to a stomatal conductance
of 37.5 mmol O
3
m
2
s
1
when the ozone concentration
at the leaf surface is 40 ppb. In this study, ux thresh-
olds are used to highlight the spatial dierences that
exist both between dierent European locations and to
provide a comparison with the total cumulative ux and
exposure-based approaches. The 1.5 nmol m
2
s
1
threshold value has also been used to analyse the beech
data, but it is acknowledged that, for beech, because of
the lower g
max
value associated with this species, this
threshold may be too high. Rather, the value is used
solely to give an indication of the spatial variation in
higher stomatal ozone uxes to both beech and wheat.
The spatial variation in AOT40 values is considerable,
with values ranging from approximately 2000 to 9000
ppb.h for wheat and 300016000 ppb.h for beech. The
AOT40 value is lowest in Spain, although ozone con-
centrations around 40 ppb did frequently occur. In
comparison, the cumulative ozone uptake over the
growing season shows much less variation between grid
squares, with a range from 6.43 to 7.35 mmol m
2
for
wheat and 6.55 to 7.03 mmol m
2
for beech. Use of the
cumulative uptake above 1.5 nmol O
3
m
2
s
1
gives a
dierent spatial pattern again, with a range across the
four grid squares of 2.44 to 4.48 mmol m
2
for wheat
and 0.62 to 1.46 mmol m
2
for beech. Ranking the
values using this index results in an opposite ordering of
the grid squares to that obtained using the AOT40
approach, with the order being the same for both spe-
cies. The highest values of TCUO
3
occurred in the
Spanish grid square. For wheat, this was a consequence
of the earlier growing season which resulted in the
high mid-day ozone concentrations occurring with con-
ditions optimal for stomatal conductance and hence
ozone uptake.
Further analysis of both the wheat and beech data
considered the mean relative g values (for PFD, tem-
perature, VPD, SMD) over the 6 h periods. The corre-
sponding stomatal conductance to ozone (GO
3
) and
ozone concentrations (O
3
) were also plotted to give an
indication of the interaction between the environmental
conditions, and the ozone concentrations over the diur-
nal period. Fig. 4 shows an example of this analysis
made for beech located in the Czech Republic grid
square during July. From Fig. 4, it is evident that, dur-
ing the daylight period when ozone concentrations were
highest, stomatal conductance tended to be at its lowest.
For the 12 hour period, VPD, and, to a lesser extent,
temperature, were the most important factors limiting g
s
and hence ozone uptake. In the early evening, when
ozone concentrations are still high, VPD remains the
predominant limiting factor to ozone uptake, although
the decreasing levels of irradiance also signicantly
reduce g
s
. In contrast, during the early morning period
the environmental conditions are at their optimum for
Fig. 3. Mean monthly stomatal ozone ux (nmol O
3
m
2
s
1
) to beech
in June during 1994.
Fig. 2. Mean monthly stomatal ozone ux (nmol O
3
m
2
s
1
) to
wheat in June during 1994.
Fig. 1. Distribution of AOT40 values (ppb-h) during June in 1994.
L.D. Emberson et al. / Environmental Pollution 109 (2000) 403413 409
conductance with only a small limitation to g
s
exerted
by low temperatures. As such, it is during this period,
when ozone concentrations are actually at their lowest,
that the highest ozone uptake rates occur. This pattern
was found for all locations, although the limitations due
to environmental factors were greatest in the Czech
Republic grid square, where the ozone concentrations
tended to be highest.
The role that VPD plays in limiting ozone uptake was
further substantiated in separate analyses conducted for
both beech and wheat. This involved identication of
those periods when ozone concentrations exceeded 60
ppb. For all situations, and for both species, it was clear
that high VPDs were the main environmental variable
limiting g
s
, and hence ozone ux, during these episodes
of high ozone concentrations. SMD did not cause any
signicant limitation to wheat stomatal ozone ux since
the wheat-growing season was completed before SMD
had accumulated to a high enough level to limit stoma-
tal conductance. However, an eect of SMD in limiting
stomatal ux to beech was observed later in the growing
season in Spain.
5. Conclusions
A new deposition model has been presented, which is
intended to allow the calculation of stomatal ozone
uptake for major vegetation types across Europe. The
model was designed to link to European-scale photo-
oxidant models, enabling estimation of stomatal uxes
over the whole of Europe and for multi-annual time-
periods, in order to improve future damage estimates
for vegetation, and hence improve the data available
for policy analysis. The ux model takes into account
the major factors believed to inuence stomatal uptake
of gases by vegetation, notably phenology, irradiance,
temperature, VPD and SMD.
The results of this initial modelling study have
demonstrated the signicant dierences in spatial pat-
terns across Europe that result from using a ux, rather
than an exposure, based approach. This clearly indi-
cates the limitations of using AOT40, and its exceed-
ance, as the basis for any assessment of the relative risk
Table 2
Calculated total (CUO
3
) and threshold (TCUO
3
) cumulative ozone uptake values for beech and wheat over the 1994 growing season compared to
equivalent values of AOT40
Location CUO
3
(mmol m
2
) TCUO
3
(mmol m
2
) AOT40
a
(ppb.h)
Wheat
Sweden 7.35 3.33 4714
UK 6.54 3.23 4407
Czech Republic 6.43 2.44 8730
Spain 6.65 4.48 2283
Beech
Sweden 6.55 1.24 4514
UK 6.68 0.96 4896
Czech Republic 7.03 0.62 15249
Spain 6.93 1.46 3856
a
AOT 40 is calculated over the modelled growing season and not for the xed 3- or 6-month period currently used for Level I evaluations for
agricultural crops and forest trees, respectively.
Fig. 4. Mean g relationships (with g
pot
equal to 1) and relationship
between O
3
concentrations (ppb) and GO
3
(mmol O
3
m
2
s
1
) for
beech during July in the Czech Republic grid square.
410 L.D. Emberson et al. / Environmental Pollution 109 (2000) 403413
of plant damage across Europe. If AOT40 had been
used as a basis for assessing risk, then clearly the highest
values would be found in central Europe; in contrast,
values of the ux indices in central Europe were either
comparable to, or lower than, those in the other grid
squares.
In this study, we have expressed ux both in terms of
total cumulative uptake over the growing season
(CUO
3
), and the cumulative uptake above a threshold
ux of 1.5 mmol m
2
s
1
(TCUO
3
). These two indices
produced very dierent patterns in the four grid squares
investigated, and it is clear that more work is needed to
determine whether such a ux threshold exists and, if so,
what value would be appropriate to replace the empiri-
cally derived 40 ppb value used in the AOT40 index.
Since plants are known to possess a defence capacity
(e.g. through antioxidant activity), and have a capacity
to repair injured tissue, it is to be expected that there
will be some threshold level of ozone ux. Weber et al.
(1993), for example, dene a `threshold ozone uptake
rate', at which ozone ux exceeds the defence response.
However, in one of the rst empirical analyses of
the size of the ux threshold (Pleijel et al., 1999), a
series of open-top chamber studies with cereals found
that the total ux provided a better t to the experi-
mental data than either the AOT40 values or cumulative
uptake above any critical ux threshold. The situation is
complicated by the variable size of the defence capacity,
and the fact that this can be both induced by ozone
exposure, or exhausted by long-term exposure to high
ozone concentrations (Musselman and Massman, 1999).
In terms of specic factors determining ozone ux,
phenology was highly signicant under our model
assumptions, since the timing of the growing season
determines both the ozone concentrations and the
environmental conditions experienced by the plant, as
was shown particularly for the results in the Spanish
grid square. The workshops at which critical level values
have been xed have recommended that, for crop
species, AOT40 should be calculated over the period
most relevant for the particular crop and region
(Ka renlampi and Ska rby, 1996). However, for the
purposes of assessing the eects of emission control
strategies on AOT40 exceedance, modelling has, for
practical purposes, been carried out using xed time
periods across Europe. It is clear that the use of ux,
rather than exposure, indices will further complicate the
process of assessing risk to dierent target species across
Europe. This is a consequence of the need to consider
the temporal variation, not only in ozone concen-
trations, but also in environmental variables aecting
both phenology and stomatal conductance.
This study suggests that, with the model formulation
employed, the most important environmental variable
was VPD, since high VPDs that limit g
s
tend to occur
with high ozone concentrations. SMD was also an
important limiting factor for beech, but not wheat, in
the Spanish grid square since the length of the wheat-
growing season was too short to allow the build up of
SMDs large enough to signicantly limit g
s
.
There are three obvious limitations to this study. The
rst regards the quality and resolution of the input data.
The assumptions required to estimate leaf temperature
and VPD from the modelled dataset are particularly
important. The 150 km150 km spatial scale means
that local variations in climate, such as those caused by
increasing elevation, will not be simulated by the map-
ping procedure. In addition, the use of 6-hourly values
means that the more subtle co-variations of ozone con-
ductance and ozone concentration, which occur over the
course of a day, cannot be simulated.
Secondly, the g
s
dataset, in terms of the species-specic
parameters and relationships with environmental vari-
ables, is based on limited information, and it does not
include the wide variation in g
s
responses which may
exist within a species across Europe. For these reasons,
our results should be viewed as indicative rather than
predictive at this stage. Some preliminary comparisons
have been made of our model predictions with both eld
and chamber measurements on Picea abies (Emberson et
al., 2000; Karlsson et al., 2000). These indicate a reason-
able correlation between observed and predicted values,
although, as expected, the precise parameterisation at
a particular site diered from the generalised para-
meterisation used in our model. However, they also
demonstrate some areas in which the model formu-
lation could be improved, such as interactive eects
of environmental parameters (e.g. SMD and VPD)
and long-term eects of ozone exposure on stomatal
responses. It is clear that further work is needed to
validate and modify the stomatal component of our
model.
Thirdly, the omission of a component calculating
deposition to external surfaces in the model means that
the calculated uxes are likely to systematically over-
estimate the ux to an individual leaf at the top of the
canopy.
An improved ozone deposition module is currently
being developed which can be linked to the stomatal
ux model presented here. This will provide a better
estimate of the bulk canopy resistance and hence the
ozone concentration gradient. It is intended that this
deposition module will be incorporated into the 5050
km
2
Eulerian EMEP ozone model, both to provide
improved deposition and ux estimates, and to examine
the eects of variable ozone deposition on atmospheric
ozone concentrations. The development of a consistent
methodology to model total deposition and absorbed
stomatal dose should provide a further improved esti-
mate of ozone risk, especially in cases where there may
be large spatial variation, e.g. between irrigated and
non-irrigated areas within a grid square.
L.D. Emberson et al. / Environmental Pollution 109 (2000) 403413 411
Acknowledgements
This project was supported by the UK Department of
the Environment, Transport and the Regions (DETR),
the Nordic Council of Ministers (NMR), and the Co-
operative Programme for Monitoring and Evaluation of
the Long-Range Transmission of Air Pollutants in Eur-
ope (EMEP).
References
Amiro, B.D., Gillespie, T.J., Thurtell, G.W., 1984. Injury response of
Phaseolus vulgaris to ozone ux density. Atmospheric Environment
18, 12071215.
Araus, J.L., Tapia, L., 1987. Photosynthetic gas exchange character-
istics of wheat ag leaf blades and sheaths during grain lling. Plant
Physiology 85, 667673.
Araus, J.L., Tapia, L., Alegre, L., 1989. The eect of changing sowing
date on leaf structure and gas exchange characteristics of wheat ag
leaves grown under Mediterranean climate conditions. Journal of
Experimental Botany 40, 639646.
Baldocchi, D.D., Hicks, B.B., Camara, P., 1987. A canopy stomatal
resistance model for gaseous deposition to vegetated surfaces.
Atmospheric Environment 21, 91101.
Bull, K.R., 1991. The critical loads/levels approach to gaseous pollu-
tant emission control. Environmental Pollution 69, 105123.
Campbell, G.S., 1985. Soil Physics with Basic. Transport Models for
Soil-Plant Systems. Developments in Soil Science 14. Elsevier,
Oxford.
Cramer, W., wolfgang.cramer@pik-potsdam.de.
Eamus, D., Murray, M., 1991. Photosynthetic and stomatal con-
ductance responses of Norway spruce and beech to ozone, acid mist
and frost a conceptual model. Environmental Pollution 72, 2345.
Emberson, L.D., 1997. Dening and Mapping Relative Potential Sen-
sitivity of European Vegetation to ozone. Ph.D. thesis, Imperial
College, University of London.
Emberson, L.D., Wieser, G., Ashmore, M.R., 2000. Modelling of sto-
matal conductance and ozone ux of Norway spruce: comparison
with eld data. Environmental Pollution 109, 393402.
Erisman, J.W., van Pul, A., Wyers, P., 1994. Parameterization of sur-
face resistance for the quantication of atmospheric deposition of
acidifying pollutans and ozone. Atmospheric Environment 28,
25952607.
ESA, 1992. Remote Sensing Map of Europe (1:6,000,000). European
Space Agency/ESTEC, Noordijk.
Eurostat, 1994. Crop Production statistics. Oce for Ocial Publica-
tions of the European Communities, Brussels.
FAO-Cartographia, 1980. Land Use Map of Europe (1:2,500,000).
Cartographia, Budapest.
FAO-UNESCO, 1981. Soil Map of the World, Volume V, Europe.
UNESCO, Paris.
Fuhrer, J., Ska rby, L., Ashmore, M.R., 1997. Critical levels for ozone
eects on vegetation in Europe. Environmental Pollution 97, 91106.
Fuhrer, J., Grandjean Grimm, A., Tschannen, W., Shariat-Madari,
H., 1992. The response of spring wheat (Triticum aestivum L.) to
ozone at higher elevations. II Changes in yield, yield components
and grain quality in response to ozone ux. New Phytologist 121,
211219.
Gollan, T., Passioura, J.B., Munns, R., 1986. Soil water status eects
the stomatal conductance of fully turgid wheat and sunower
leaves. Australian Journal of Plant Physiology 13, 459464.
Grandjean Grimm, A., Fuhrer, J., 1992a. The response of spring
wheat (Triticum aestivum L.) to ozone at higher elevations. I Mea-
surement of ozone and carbon dioxide uxes in open-top eld
chambers. New Phytologist 121, 201210.
Grandjean Grimm, A., Fuhrer, J., 1992b. The response of spring
wheat (Triticum aestivum L.) to ozone at higher elevations. III
Responses of leaf canopy gas exchange, and chlorophyll uores-
cence to ozone ux. New Phytologist 122, 321328.
Gru nhage, L., Ja ger, H.J., 1994. Inuence of atmospheric conductivity
on the ozone exposure of plants under ambient conditions: con-
siderations for establishing ozone standards to protect vegetation.
Environmental Pollution 85, 125129.
Gru nhage, L., Haenel, H.-D., 1997. PLATIN (PlantAtmosphere
Interaction) I: a model of plantatmosphere interaction for esti-
mating absorbed doses of gaseous air pollutants. Environmental
Pollution 98, 3750.
Gru nhage, L., Ja ger, H.-J., Haenel, H.-D., Hanewald, K., Krupa, S.,
1997. PLATIN (PlantAtmosphere Interaction) II: co-occurrence of
high ambient ozone concentrations and factors limiting plant
absorbed dose. Environmental Pollution 98, 5160.
Gru ters, U., Fangmeier, A., Ja ger, H.-J., 1995. Modelling stomatal
responses of spring wheat (TriticumaestivumL. cv. Turbo) to ozone at
dierent levels of water supply. Environmental Pollution 87, 141149.
Hosker Jr., R.P., 1986. Practical application of air pollutant deposi-
tion models current status, data requirements and research needs.
In: Legge, A.H., Krupa, S.V. (Eds.), Air Pollutants and their Eects
on the Terrestrial Ecosystem. John Wiley, New York, pp. 505567.
Jackson, R.B., Canadell, J., Ehleringer, J.R., Mooney, H.A., Sala,
O.E., Schulze, E.D., 1996. A global analysis of root distributions for
terrestrial biomes. Oecologia 108, 389411.
Jakobsen, H.A., Jonson, J.E., Berge, E., 1996. Transport and Deposi-
tion Calculations of Sulphur and Nitrogen Compounds in Europe
for 1992 in the 50 km Grid by Use of the Multi-layer Eulerian
Model (EMEP/MSC-W Report 2/96). The Norwegian Meteor-
ological Institute, Oslo, Norway.
Jarvis, P.G., 1976. The interpretation of the variations in leaf water
potential and stomatal conductance found in canopies in the eld.
Philosophical Transactions of the Royal Society, London B 273,
593610.
Jones, H.G., 1992. Plants and Microclimate: A Quantitative Approach
to Environmental Plant Physiology, 2nd Edition. Cambridge Uni-
versity Press, Cambridge.
Ka renlampi, L., Ska rby, L. (Eds.), 1996. Critical Levels for Ozone in
Europe: Testing and Finalizing the Concepts (UN/ECE Workshop
Report). University of Kuopio, Dept. of Ecology and Environ-
mental Science, Kuopio, Finland.
Karlsson, P.E., Pleijel, H., Pihl Karlsson, G., Medin, E.L., Ska rby, L.,
2000. Simulations of stomatal conductance and ozone uptake to
Norway spruce saplings in open-top chambers. Environmental Pol-
lution 109, 441449.
Kerstiens, G., 1995. Cuticular water performance of European trees
and shrubs grown in polluted and unpolluted atmospheres, and its
relation to stomatal response to humidity in beech (Fagus sylvatica
L.). New Phytologist 112, 1319.
Ko rner, C., 1994. Leaf diusive conductances in the major vegetation
types of the globe. In: Schulze, E.D., Caldwell, M.M. (Eds.), Eco-
physiology of Photosynthesis. Ecological Studies Vol. 100. Springer,
Heidelberg, pp. 463490.
Ko rner, C., Peterer, J., Altrichter, Ch., Meusburger, A., Slovik, S.,
Zoschg, M., 1995. A simple empirical model to estimate annual dry
deposition of atmospheric pollutants in needles of spruce and pine.
Allgemeine Forst- und Jagdzeitung 166, 19.
Kostrowski, 1984. Types of Agriculture Map of Europe. Polish
Academy of Sciences, Warsaw
Kramer, K., 1994. Selecting a model to predict the onset of growth of
Fagus sylvatica. Journal of Applied Ecology 31, 172181.
Kramer, K., 1995. Phenotypic plasticity of the phenology of seven
European tree species in relation to climatic warming. Plant, Cell
and Environment 18, 93104.
Laisk, A., Kull, O., Moldau, H., 1989. Ozone concentration in leaf
intercellular air spaces is close to zero. Plant Physiology 90, 11631167.
412 L.D. Emberson et al. / Environmental Pollution 109 (2000) 403413
Lehnherr, B., Ma chler, F., Grandjean Grimm, A., Fuhrer, J., 1988.
The regulation of photosynthesis in leaves of eld-grown spring
wheat (Triticum aestivum L. cv Albis) at dierent levels of ozone in
ambient air. Plant Physiology 88, 11151119.
Lo vblad, G., Grennfelt, P., Ka renlampi, L., Laurila, T., Mortensen,
L., Ojanpera , K., Pleijel, H., Semb, A., Simpson, D., Ska rby, L.,
Tuovinen, J.-P., Trseth, K., 1996. Ozone exposure mapping in the
Nordic countries. TemaNord 1996:522. Nordic Council of Minis-
ters, Copenhagen
Machado, E.C., Lago a, A.M.M.A., 1994. Trocas gasosas e con-
duta ncia estoma tica em tre s especie s de gram neas. Bragantia,
Campinas 53, 141149.
Mikkelsen, T.N., 1995. Physiological responses of Fagus sylvatica L.
exposed to lowlevels of ozone in open-top chambers. Trees 9, 355361.
Milthorpe, F.L., Moorby, J., 1974. An Introduction to Crop Physiol-
ogy. Cambridge University Press
Monteith, J.L., Unsworth, M., 1990. Principles of Environmental
Physics. 2nd Edition. Arnold, London.
Morgan, J.A., 1984. Interaction of water supply and N in wheat. Plant
Physiology 76, 112117.
Musselman, R.C., Massman, W.J., 1999. Ozone ux to vegetation and
its relationship to plant response and ambient air quality standards.
Atmospheric Environment 33, 6573.
Osonubi, O., Davies, W.J., 1980. The inuence of water stress on the
photosynthetic performance and stomatal behaviour of tree seed-
lings subjected to variation in temperature and irradiance. Oecolo-
gia (Berl.) 45, 310.
Pearson, M., Manseld, T.A., 1993. Interacting eects of ozone and
water stress on the stomatal resistance of beech (Fagus sylvatica L.).
New Phytologist 123, 351358.
Peterson, R.F., 1965. Wheat: Botany, Cultivation, and Utilization.
Leonard Hill Books, London.
Pleijel, H., Wallin, G., Karlsson, P.E., Ska rby, L., Sellde n, G., 1994.
Flux measurements in open-top chambers. In: Ja ger, H.J., Uns-
worth, M., De Temmerman, L., Mathy, P. (Eds.), Eects of Air
Pollution on Agricultural Crops in Europe. CEC Proceedings of the
Final Symposium of the European Open-Top Chambers Project.
Tervuren, Belgium, CEC, Brussels, pp. 193211.
Pleijel, H., Danielsson, H., Pihl Karlsson, G., Grelang, J., Karlsson,
P.E., 1999. An ozone ux-response relationship for cereals. In:
Fuhner, J., Ackermann, B. (Eds.), Critical Levels for Ozone
Level II. Environmental Documentation No. 115. Swiss Agency
for Environment, Forest and Landscape, Bern, Switzerland, pp. 95
99.
Sanders, G., Clark, A.C., Colls, J.J., 1991. The inuence of open-top
chambers on the growth and development of eld bean. New Phy-
tologist 117, 439447.
Simpson, D., 1993. Photochemical model calculations over Europe for
two extended summer periods: 1985 and 1989. Model results and
comparisons with observations. Atmospheric Environment 27A,
921943.
Simpson, D., 1995. Biogenic emissions in Europe 2: implications for
ozone control strategies. Journal of Geophysical Research 100
(D11), 2289122906.
Taylor, G., Dobson, M.C., 1989. Photosynthetic characteristics, sto-
matal responses and water relations of Fagus sylvatica: impact of air
quality at a site in southern Britain. New Phytologist 113, 265273.
Thornthwaite, C.W., Mather, J.R., 1957. Instructions and Tables for
Computing Potential Evaporation and the Water Balance. Publica-
tions in climatology X No.3, Centerton, New Jersey.
Tognetti, R., Johnson, J.D., Michelozzi, M., 1995. The response of
European beech (Fagus sylvatica L.) seedlings from two Italian
populations to drought and recovery. Trees 9, 348354.
Tuebner, F., 1985. Messung der Photosynthese und Transpiration an
Weizen, Kartoel und Sonnenblume. Diplomarbeit University
Bayreuth, West Germany
Wang, D., Hinckley, M., Cumming, A.B., Braatne, J., 1995. A com-
parison of measured and modeled ozone uptake into plant leaves.
Environmental Pollution 89, 247254.
Weber, P., Rennenberg, H., 1996. Dependency of nitrogen dioxide
(NO
2
) uxes to wheat (Triticum aestivum L.) leaves from NO
2
con-
centration, light intensity, temperature and relative humidity deter-
mined from controlled dynamic chamber experiments. Atmospheric
Environment 30, 30013009.
Weber, J.A., Clark, C.S., Hogsett, W.E., 1993. Analysis of the rela-
tionship among O
3
uptake, conductance and photosynthesis in nee-
dles of Pinus ponderosa. Tree Physiology 13, 157172.
Wesely, M.L., 1989. Parameterization of surface resistances to gaseous
deposition in regional-scale numerical models. Atmospheric Envir-
onment 23, 12931304.
L.D. Emberson et al. / Environmental Pollution 109 (2000) 403413 413