towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry markus löw...

29
Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1 , Manuela Blumenröther 2 , Karl-Heinz Häberle 1 , Wolfgang Oßwald 2 and Rainer Matyssek 1 1 Ökophysiologie der Pflanzen 2 Fachgebiet Krankheiten der Waldbäume Technische Universität München, Am Hochanger 13, 85354 Freising, Ger

Upload: oliver-arnold

Post on 04-Jan-2016

217 views

Category:

Documents


0 download

TRANSCRIPT

Page 1: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

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

Page 2: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

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

Page 3: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

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

Page 4: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

O3 flux model at „Kranzberger Forst“

Linear relationship in 2002 and 2004 = years without drought

Page 5: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

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“

Page 6: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

Assessment of gas exchange

• Light saturated rate of photosynthesis (Amax)

• Measured during CASIROZ sampling campaigns

• Diagnostic test under „steady state“ conditions

• Licor 6400

Page 7: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

Annual course of photosynthesis

Overall O3 effect on net CO2 uptake rate:

p = 0.04 (GLM, rep. meas.)

2004

means ± SDn = 4-5 trees

Page 8: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

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

Page 9: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

Annual course of photosynthesis

Unusual annual course

2003 (drought year)

means ± SDn = 4-5 trees

Page 10: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

Error bars indicate standard errorsun crown

***

ozone effect disappeared during drought

conductance was minimal in both treatments in 2003

stomatal conductance gH2O 2003 - 2004

Page 11: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

Annual course of photosynthesis

Unusual annual course

2003 (drought year)

means ± SDn = 4-5 trees

Page 12: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

Annual course of photosynthesis

„normal“ annual course

r2 = 0.57p < 0.05

2004

means ± SDn = 4-5 trees

Page 13: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

Ozone effect on photosynthesis

means ± SDn = 4-5 trees

Page 14: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

Ozone effect on photosynthesis

means ± SDn = 4-5 trees

(drought year)

Page 15: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

Ozone effect on photosynthesis

p < 0.05

senescencein October

(drought year)

Drought overrules ozone effect, linear correlation only during summer months

Page 16: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

Ozone effect on photosynthesis

means ± SDn = 4-5 trees

Ozone effect on photosynthesis

Page 17: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

Linear correlation only in summer months (June to September)

n.s.leaf expansionin May

senescencein October

Ozone effect on photosynthesis

Page 18: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

Pause

Pause

– linking leaf gas exchange with biochemistry

Page 19: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

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)...)

Page 20: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

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]

Page 21: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

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

Page 22: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

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

*

Page 23: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

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

Page 24: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

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

Page 25: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

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

Page 26: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

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

Page 27: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

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….

Page 28: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

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)

Page 29: Towards flux-based ozone risk assessment – linking leaf gas exchange with biochemistry Markus Löw 1, Manuela Blumenröther 2, Karl-Heinz Häberle 1, Wolfgang

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...