towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry markus löw...
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Towards flux-based ozone risk assessment
– linking leaf gas exchange with biochemistry
Markus Löw1, Manuela Blumenröther2, Karl-Heinz Häberle1, Wolfgang Oßwald2 and Rainer Matyssek1
1Ökophysiologie der Pflanzen2Fachgebiet Krankheiten der Waldbäume
Technische Universität München, Am Hochanger 13, 85354 Freising, Germany
O3 flux model at „Kranzberger Forst“
• Based on Emberson et al. (2000)
• Extended for drought conditions
• Validated for adult beech under site conditions
• Working (2002, 2003, 2004)
• Nunn et al. (2005) Env. Poll. 137(3): 494-506
O3 flux model at „Kranzberger Forst“
• COU: cumulative stomatal ozone uptake (AFst0)
• gmax measured individually per tree and crown position
• growing season determined individually per tree and crown position
• modelled on a 10 minute basis• soil moisture function
implemented
O3 flux model at „Kranzberger Forst“
Linear relationship in 2002 and 2004 = years without drought
End of July 2003: drought became a limiting factor
No linear relationship in 2003, so AOT40 predicts misleading high risk
O3 flux model at „Kranzberger Forst“
Assessment of gas exchange
• Light saturated rate of photosynthesis (Amax)
• Measured during CASIROZ sampling campaigns
• Diagnostic test under „steady state“ conditions
• Licor 6400
Annual course of photosynthesis
Overall O3 effect on net CO2 uptake rate:
p = 0.04 (GLM, rep. meas.)
2004
means ± SDn = 4-5 trees
Annual course of photosynthesis
No statistically significant O3 effects in 2003
20042003
*
* *
*
Significant drought effects in 2003: 1xO3 p < 0.001; 2xO3 p = 0.02
*
*
means ± SDn = 4-5 trees
Annual course of photosynthesis
Unusual annual course
2003 (drought year)
means ± SDn = 4-5 trees
Error bars indicate standard errorsun crown
***
ozone effect disappeared during drought
conductance was minimal in both treatments in 2003
stomatal conductance gH2O 2003 - 2004
Annual course of photosynthesis
Unusual annual course
2003 (drought year)
means ± SDn = 4-5 trees
Annual course of photosynthesis
„normal“ annual course
r2 = 0.57p < 0.05
2004
means ± SDn = 4-5 trees
Ozone effect on photosynthesis
means ± SDn = 4-5 trees
Ozone effect on photosynthesis
means ± SDn = 4-5 trees
(drought year)
Ozone effect on photosynthesis
p < 0.05
senescencein October
(drought year)
Drought overrules ozone effect, linear correlation only during summer months
Ozone effect on photosynthesis
means ± SDn = 4-5 trees
Ozone effect on photosynthesis
Linear correlation only in summer months (June to September)
n.s.leaf expansionin May
senescencein October
Ozone effect on photosynthesis
Pause
Pause
– linking leaf gas exchange with biochemistry
Measured parameters
• Phosphoenolpyruvate-carboxylase activity (PEPc)[nkat/mg protein] in sun and shade leaves
• Starch content [mg glucose equivalent/g dw] in sun and shade leaves
CO2-fixing enzyme, contributing to anaplerotic pathways, stress induced
Ambivalent results for O3 effects on carbohydrates
Activity is said to increase under O3/stress impact(Luethy-Krause et al. 1990 (Norway spruce, Scots pine);Landolt et al. 1994, 1997 (poplar, birch); Saurer et al. 1995 (birch); Fontaine 1999 (Aleppo pine); Lütz et al. 2000 (beech); Inclam et al. 2005 (Aleppo pine)...)
0.0
0.2
0.4
0.6
0.8
1.0
1.2
1.4
May. 03 Jun. 03 Jul. 03 Aug. 03 Sep. 03 Oct. 03
date
[nk
at
mg
pro
tein
-1]
Specific PEPc activity 2003
1xO3 sun 2003 2xO3 sun 2003 1xO3 shade 2003 2xO3 shade 2003
No significant annual effect of O3 on PEPc in sun & shade leaves 2003
mv ± se
0.0
0.2
0.4
0.6
0.8
1.0
1.2
1.4
May. 03 Jun. 03 Jul. 03 Aug. 03 Sep. 03 Oct. 03
date
[nk
at
mg
pro
tein
-1]
Significant correlations to COU in sun & shade leaves 2003
But activity in 2xO3 leaves always lower at any given flux
R2 = 0.9619 R2 = 0.9596
R2 = 0.8195
R2 = 0.9945
0.0
0.2
0.4
0.6
0.8
1.0
1.2
1.4
0 5 10 15 20 25 30 35 40
COU [mmol m-2]
[nk
at
mg
pro
tein
-1]
R2 = 0.9619 R2 = 0.9596
R2 = 0.8195
R2 = 0.9945
0.0
0.2
0.4
0.6
0.8
1.0
1.2
1.4
0 5 10 15 20 25 30 35 40
COU [mmol m-2]
[nk
at
mg
pro
tein
-1]
Specific PEPc activity vs. COU 2003 (dry year!)
p < 0.05each
0.0
0.2
0.4
0.6
0.8
1.0
1.2
1.4
May. 04 Jun. 04 Jul. 04 Aug. 04 Sep. 04 Oct. 04
date
[nk
at
mg
pro
tein
-1]
0.0
0.2
0.4
0.6
0.8
1.0
1.2
1.4
May. 04 Jun. 04 Jul. 04 Aug. 04 Sep. 04 Oct. 04
date
[nk
at
mg
pro
tein
-1]
Specific PEPc activity 2004
1xO3 sun 2004 2xO3 sun 2004 1xO3 sha 2004 2xO3 sha 2004
p = 0.038
Significantly decreased PEPc activity in 2xO3 sun leaves throughout 2004
mv ± se
*
Rising activity in shade leaves is comparable to the pattern in 2003, but is not significantly
correlated to COU
2xO3 sun leaves are not responding to COU in 2004; 1xO3 sun leaves are significantly negative
correlated to flux
R2 = 0.0881
R2 = 0.9341
R2 = 0.626
R2 = 0.6573
0.0
0.2
0.4
0.6
0.8
1.0
1.2
1.4
0 10 20 30 40
COU [mmol m-2]
sp
ec
. ac
tiv
ity
[n
ka
t m
g p
rote
in-1
]
R2 = 0.0881
R2 = 0.9341
R2 = 0.626
R2 = 0.6573
0.0
0.2
0.4
0.6
0.8
1.0
1.2
1.4
0 10 20 30 40
COU [mmol m-2]
sp
ec
. ac
tiv
ity
[n
ka
t m
g p
rote
in-1
]
Specific PEPc activity vs. COU 2004
p < 0.01
Chronic effects of O3 result in decreased PEPc activity
0
5
10
15
20
25
May. 03 Jun. 03 Jul. 03 Aug. 03 Sep. 03 Oct. 03
date
[mg
glc
eq
uiv
ale
nt
g d
w-1
]
0
5
10
15
20
25
May. 03 Jun. 03 Jul. 03 Aug. 03 Sep. 03 Oct. 03
date
[mg
glc
eq
uiv
ale
nt
g d
w-1
]
Starch concentrations in leaves 2003 (dry year)
1xO3 sun 2003 2xO3 sun 2003 1xO3 shade 2003 2xO3 shade 2003
Significantly decreased starch contents in 2xO3 sun leaves throughout May, June & July
p = 0.023
mv ± se
*allocationprocesses
0
5
10
15
20
25
May. 04 Jun. 04 Jul. 04 Aug. 04 Sep. 04 Oct. 04
date
[mg
glc
eq
uiv
ale
nt
g d
w-1
]
Starch concentrations in leaves 2004
1xO3 sun 2004 2xO3 sun 2004 1xO3 sha 2004 2xO3 sha 2004
Higher starch levels in 2004Significantly reduced starch contents in 2xO3
sun
p = 0.009
Carbohydrates used for repair/detoxification processes in 2xO3 sun leaves?
Less photosynthetic products available under 2xO3?
**allocationprocesses
mv ± se
0
20
40
60
80
100
0 5 10 15 20 25 30COU [mmol m-2]
rela
tiv
e s
tarc
h, A
ma
x [
%]
0
20
40
60
80
100
0 5 10 15 20 25 30COU [mmol m-2]
rela
tiv
e s
tarc
h, A
ma
x [
%]
1xO3 = 100%
0
20
40
60
80
100
0 5 10 15 20 25 30COU [mmol m-2]
rela
tiv
e s
tarc
h, A
max
[%
]
Relative starch and Amax vs. COU in sun leaves 2004
r2 = 0.738p < 0.05
Annual response patterns of Amax and starch to O3 are significantly correlated
starch Amax
Conclusions I• O3 flux model for adult beech trees in
„Kranzberger Forst“ validated & ready to use
• Discrepancy between O3 exposure and uptake especially in dry year
• Significant O3 effect on photosynthesis only in 2004
• Significantly reduced starch concentrations in sun leaves 2003 and 2004; differences between O3 regimes more pronounced in 2004
• Significant correlation of PEPc activity and COU
• Biochemical parameters indicate a chronic O3 effect
• Controversial results for PEPc indicate the need for more studies on adult trees….
Conclusions II• Drought overrules O3 impact in 2003
• Different parameters (starch + Amax) show similar responses to O3
• Risk assessment does profit from the combination of several parameters from different scaling levels
• Thus, these parameters should be included in Flux modelling (Flux + defence)
Acknowledgements
The present study is part of the Project "CASIROZ – The carbon sink strength of beech in a changing environment: Experimental risk assessment by mitigation of chronic ozone impact", which is supported by European Commission - Research Directorate-General, Environment Programme, "Natural Resources Management and Services" (EVK2-2002-00165, Ecosystem Vulnerability).
Pierre Dizengremel, UHP Nancy, France
Frank FleischmannTina Schmidt
Angela NunnThorsten Grams
Thomas Feuerbach...