Benefits of Adding Physiological Measurements to Long-term Monitoring Efforts Calcium Depletion and...

Benefits of Adding Physiological Measurements to Long-term

Monitoring Efforts

Calcium Depletion and Red Spruce / Sugar Maple Decline as Examples

USDA Forest Service

Paul SchabergKelly BaggettJohn BenninkBrynne Lazarus Paula Murakami

The University of Vermont

Rubenstein School, Burlington, VT

Donald DeHayesGary HawleyCathy BorerTammy CoeMichelle TurnerBrett HuggettJosh Halmanand many students…

Winter Injury of Red Spruce

Cold tolerance of several north temperate conifers(sampled in Wolcott, VT on March 3, 1998)

Species Mean cold tolerance (ºC)

Red spruce

White pine

-61.0

-90.0

-38.1

-59.5

Eastern hemlock

White spruce

Red pine -90.0

DeHayes et al. 2001

Mechanisms for acid-induced

freezing injury of red spruceAcid deposition (H+)Acid deposition (H+)

Foliar Ca leaching(2-8X increase, P<0.05)

H+ displacementof membrane-associated Ca

(-20%, P<0.05)

Membrane destabilization(+20%, P<0.05)

& reduction in messenger Ca

Reduced cold tolerance(-10%, P<0.05)

Increased susceptibilityto freezing injury

DeHayes et al. 1999, Schaberg et al. 2000, Schaberg et al. 2001

Unresolved issues…

•Relevance to field?•Soil Ca depletion?•Just cold tolerance?•Just red spruce?

Providing a Broader Context

Field verificationSoil-based Ca depletionStress response systems Other tree species

2003 – high winter injury year

Historic Context of 2003 Winter Injury

0

10

20

30

40

50

60

70

80

90

100

1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004

Year

Mea

n %

cur

rent

-yea

r fo

liage

inju

red

Colebrook, NH

Mt. Moosilauke, NH

Whiteface Mtn., NY

2003 sampling: 185 plots at 28 sites in VT,

NH, eastern NY, and Northwestern MA Lazarus et al. 2004

2003 sampling: 185 plots at 28 sites in VT,

NH, eastern NY, and Northwestern MA

96% dominant and codominant trees effected

65% current-year foliage lost

56% buds killed

Lazarus et al. 2004

Statistical spatial analysis -significantly greater injury at:

• Higher elevations

• Western longitudes

• West-facing plots

Lazarus et al. 2006

HBEF watershed 1 (wollastonite) HBEF watershed 1 (wollastonite) compared to watershed 6 (reference) compared to watershed 6 (reference)

Wollastonite Treated Oct. 1999 (CaSiO3)

Reference

Red Spruce Winter injury

Hawley et al. 2006

Slide 15\

0

10

20

30

40

50

60

70

80

90

All Dominant and codominant Intermediate, suppressed &

Tree crown class

Win

ter

inju

ry o

f cu

rren

t-yea

r fo

liage

(%) Reference

Ca-addition

*

**

ns

understory

Environmental Change/Stress SignalTemperature extremes, fungal elicitors, wounding, drought, ozone, etc.

Plant Adaptation to Environmental Change/StressAlterations in cold tolerance, stomatal conductance,

antimicrobial defense, carbon allocation, etc.

vacuole

protein

Ca2+

enzyme activity, genetictranscription, etc.

changes in cell physiology

Ca2+

Ca2+ Ca2+

Cytoplasmic Ca binds to specific proteins (e.g.

calmodulin).

cell membrane

cell wall

Signal transduction: Camoves from area of high concentration (e.g., extracellular mCa or organelle stores) to the low Ca cytosol.

Our Overarching Hypothesis

Acid Rain, Calcium Depletionand

Immune Dysfunction

HBEF watershed 1 (wollastonite) HBEF watershed 1 (wollastonite) compared to watershed 6 (reference) compared to watershed 6 (reference)

Wollastonite Treated Oct. 1999 (CaSiO3)

Reference

Red Spruce Winter injury

Hawley et al. in press

Slide 15\

0

10

20

30

40

50

60

70

80

90

All Dominant and codominant Intermediate, suppressed &

Tree crown class

Win

ter

inju

ry o

f cu

rren

t-ye

ar f

olia

ge (

%)

ReferenceCa-addition

*

**

ns

understory

Red Spruce Foliar Cold Tolerance

-60

-50

-40-30

-20

-10

0

Ca addition Reference

Hubbard Brook Watershed

Co

ld T

ole

ran

ce (o

C)

* P = 0.0431

Foliar Cold Tolerance

Halman et al. in prep.

Carbohydrate MetabolismCarbohydrate Metabolism

Cell energy relationsCell energy relations Sugars important to cold toleranceSugars important to cold tolerance Osmotic control during freezingOsmotic control during freezing Act as an “antifreeze” agentAct as an “antifreeze” agent Direct cryoprotection of biomoleculesDirect cryoprotection of biomolecules Influenced by Ca deficiencyInfluenced by Ca deficiency

Foliar CarbohydratesFoliar Carbohydrates

Total Sugars - 2005 Foliage

40

4550

5560

65

7075

8085

90

Nov. 2005 Feb. 2006

Su

gar

Co

nce

ntr

atio

n (

mg

/g)

Reference

Ca-add

Fig. 2.Fig. 2. Total sugar levels in current-year foliage were significantly Total sugar levels in current-year foliage were significantly greater in WS1 than in WS6 in both fall and winter collections; * = (p< 0.05)greater in WS1 than in WS6 in both fall and winter collections; * = (p< 0.05)

*

**

*

Individual SugarsIndividual SugarsTable 1.Table 1. Individual sugar concentrations (mean ± SE, n = 30) in current-year foliage Individual sugar concentrations (mean ± SE, n = 30) in current-year foliage(mg g(mg g-1-1). Three major sugars tested had significantly greater means between ). Three major sugars tested had significantly greater means between watersheds from the fall collection. Data collected from winter collection watersheds from the fall collection. Data collected from winter collection showed significant differences between watersheds in sucrose and glucose.showed significant differences between watersheds in sucrose and glucose.

Sugar November February

Reference Ca-addition Reference Ca-addition

Fructose 24.56 ± 0.76** 27.75 ± 0.73** 25.74 ± 1.20 28.01 ± 1.18

Sucrose 4.14 ± 0.76 5.30 ± 0.74 2.83 ± 0.42** 4.00 ± 0.42**

Glucose 39.62 ± 0.83** 43.47 ± 0.78** 39.85 ± 1.48* 44.18 ± 1.42*

Stachyose 0.232 ± 0.03* 0.308 ± 0.03* 3.41 ± 0.19 3.56 ± 0.19

Note: * = p<0.1; ** = p<0.05Note: * = p<0.1; ** = p<0.05

Antioxidant EnzymesAntioxidant Enzymes Enzymes responsible for scavenging Enzymes responsible for scavenging

harmful reactive oxygen speciesharmful reactive oxygen species

Inadequate function can result in Inadequate function can result in cellular and membrane dysfunction, cellular and membrane dysfunction, and/or cell deathand/or cell death

Assaying ascorbate peroxidase (APX) – Assaying ascorbate peroxidase (APX) – activity dependent upon Ca availabilityactivity dependent upon Ca availability

Seasonal APX Activity

0

5

10

15

20

25

30

Nov. 2005 Feb. 2006

AP

X A

cti

vit

y (

µm

ole

s a

sc

orb

ate

m

in-1

mg

-1 p

rote

in)

Ca-add

Reference

Fig. 4.Fig. 4. APX activity (mean ± SE, n = 30) from current-year foliage. Winter APX activity APX activity (mean ± SE, n = 30) from current-year foliage. Winter APX activity is elevated in Ca-addition watershed yet relatively unchanged in reference. * = p < 0.1is elevated in Ca-addition watershed yet relatively unchanged in reference. * = p < 0.1

*

*

Field Examples Ca Depletion and Tree Decline

• Red Spruce Winter Injury

• Sugar Maple Decline

• Flowering Dogwoods and Anthracnose

• Norway spruce freezing injury

• Hemlock Woolly Adelgid Infestation

Sugar Maple Decline

• Nutritional predisposition (cations – Ca, Mg)

• Disproportionate decline following secondary stress

Mechanism of Maple Decline?

Soil Ca depletion

mCa disruption

Secondary stressesdroughtinsects disease

pollutantsetc.

Maple decline

Impairment ofstress response

system

NAMP and HHS Plots

Predicted HighCa (6)

Predicted LowCa (8)

6 sugar maples per site6 sugar maples per site84 total trees84 total trees

Till Source Model: Source EnvelopeGlacial

Movement

32 km60°

Designated site

-Courtesy of Scott Bailey et al.

• 26 HHS & NAMP sites were selected for TSM predictions. • Latitudinal and longitudinal coordinates were used to generate a source envelope.

• 8 sites predicted to contain low levels of Ca, 6 were predicted to have high levels of Ca.

0

200

400

600

800

1000

TSM Ca classes

Soil

Ca

(mg

· kg

-1 )

0

4000

8000

12000

16000

Folia

r C

a (m

g · k

g-1

)

High Low

Foliar Ca

Soil Caaa

b

b

Schaberg et al., 2006

Schaberg et al 2006

0

5

10

15

20

25

Published foliar range

% B

ran

ch d

ieb

ack

Low Moderate

Ca (P = 0.021)

Mg (P = 0.052)

Al (P = 0.035)

a ba

a bb

Schaberg et al., 2006

Schaberg et al 2006

0

40

80

120

160

0 5000 10000 15000 20000

Foliar Ca (mg · kg-1)

Bas

al g

row

th (

cm2 )

r = 0.53, P = 0.050

Schaberg et al., 2006Schaberg et al 2006

• NuPert PlotsNuPert Plots • 12 (45m x 45m) plots12 (45m x 45m) plots

• Control plots – marginal Ca nutritionControl plots – marginal Ca nutrition• Randomized Ca or Al additions – begun 1995Randomized Ca or Al additions – begun 1995

• Sugar Maple Dominant SpeciesSugar Maple Dominant Species

Hubbard Brook Experimental Hubbard Brook Experimental ForestForest

Total Foliar Cation AnalysisTotal Foliar Cation Analysis

Treatment means with different letters are significantly different (Treatment means with different letters are significantly different (PP ≤ ≤ 0.05)0.05)

based on orthogonal contrasts: Ca vs. Al and Control; Al vs. Controlbased on orthogonal contrasts: Ca vs. Al and Control; Al vs. Control

Foliar Calcium Levels

b

a

b

0

2000

4000

6000

8000

10000

Calcium Control Aluminum

Treatment

Me

an

fo

liar

Ca

(m

g •

kg

-1)

(±S

E) Minimum level for

healthy sugar maple Kolb and McCormick (1993)

Crown Health EvaluationCrown Health Evaluation

increased

dieback

decreased dieback

Percent Branch Dieback

0

1

2

3

4

5

6

7

Calcium Control Aluminum

Treatment

Me

an

% b

ran

ch

die

ba

ck

ra

tin

g (

±SE

)X2 = 6.92, p = 0.0314

Tree Increment GrowthTree Increment Growth

a

aa

a

a

a

a

b

bb

b

b

bb

b

bb

b

bb

b

0.00

1.00

2.00

3.00

4.00

5.00

1995 1996 1997 1998 1999 2000 2001 2002 2003 2004

Year

Ba

sa

l Are

a In

cre

me

nt

(BA

I) R

ati

o

Ca

Control

Al

Treatments Began

Tree Increment GrowthTree Increment Growth

a

aa

a

a

a

a

b

bb

b

b

bb

b

bb

b

bb

b

0.00

1.00

2.00

3.00

4.00

5.00

1995 1996 1997 1998 1999 2000 2001 2002 2003 2004

Year

Ba

sa

l Are

a In

cre

me

nt

(BA

I) R

ati

o

Ca

Control

Al

Treatments Began

Ice Storm

Wound Closure Amount ResultsWound Closure Amount Results

Treatment means for wound closure amounts

(mm ± SE) P-values

Ca vs. Control

Ca addition Control Al addition Control and Al vs. Al

WoundClosureAmount

0.615±0.313 -0.303±0.26 0.060 ±0.25 0.0414 0.3598

• Differences associated w/ Ca fertilization were most pronounced in wounds lower of the tree stem (100cm-130cm, P = 0.0016)

Huggett et al. submitted to CJFR

Sugar Maple Shoot Cold Tolerance

-35

-34

-33

-32

-31

-30

-29

-28

-27

-26

-25

High Low

Site soil Ca

Cold

tole

ran

ce (

oC

)

P = 0.0076***

*

March 8, 2006

Co

ld T

ole

ran

ce (C

)

Sugar Maple Shoot Cold Tolerance

Site Soil Ca Status

Synergies: Physiology and Monitoring

Gain information about the relevance of experimental evidence to trees/forests in the “real world”

Physiological measurements can add a mechanistic understanding to changes in forest health and

productivity noted via monitoring

Synergies: Physiology and Monitoring

Questions?

Benefits of Adding Physiological Measurements to Long-term Monitoring Efforts Calcium Depletion and...

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