introduction stomatal conductance regulates the rates of several necessary processes in plants...

1
Introduction Stomatal conductance regulates the rates of several necessary processes in plants including transpiration, carbon dioxide assimilation, and respiration. Both steady state and dynamic porometers are currently used to measure stomatal conductance. However, few tests or measurements have been performed to compare the results obtained with the various porometers. It is important for scientists to understand the performance of each porometer in order to compare stomatal conductance data gathered from various models. In this study, we focused on three commercially available porometers: the AP-4 (Delta-T Devices, Cambridge, UK), the SC-1 (Decagon Devices, Pullman, WA), and the LI-1600 (Li-Cor, Lincoln, NE). The objective of this study was to understand the relationship between the stomatal conductance data gathered with the different instruments An Inter-comparison of Three Commercial An Inter-comparison of Three Commercial Porometers Porometers L. Bissey 1 , D. Cobos 2 , C. Campbell 2 1. School of Earth and Environmental Science, Washington State University, 2. Decagon Devices, Inc. Figure 2. Example data showing leaf-level stomatal conductance variability. Numbers in black, blue, and purple are conductance data obtained using the AP-4, SC-1, and LI-1600 respectively. Data from different instruments were collected at different times and should not be compared among porometers. 962, 673, 410 678, 674, 362 701, 560, 383 690, 918, 425 753, 573, 322 1053, 964, 356 We measured stomatal conductance of approximately 200 leaves of various species with an AP-4, SC- 1, and LI-1600 under field conditions. Changes in environmental conditions were minimized by taking measurements with the three porometers on similar locations of each leaf in rapid succession. On each leaf, stomatal conductance measurements with the three porometers were conducted in random order to prevent systematic bias from previous porometer measurements. Each porometer was factory calibrated within 60 days of the beginning of measurements. This methodology assumes that: 1) stomatal conductance was measured on the exact same footprint on each leaf, and 2) environmental factors (radiation, temperature) didn’t change over the 1-2 minute timescale needed to make the three measurements on each leaf. 1 2 3 4 5 6 7 8 1 2 3 4 5 6 7 8 Ln LI-1600 Stom atalConductance (m mol/m 2 s) SC-1 Stom atal Conductance(m m ol/m 2 s) 1 2 3 4 5 6 7 8 1 2 3 4 5 6 7 8 Ln LI-1600 Stom atalConductance (m mol/m 2 s) LN AP-4Stom atal Conductance(m m ol/m 2 s) 1 2 3 4 5 6 7 8 1 2 3 4 5 6 7 8 LN A P-4 Stom atalConductance (m mol/m 2 s) LN SC-1Stom atal Conductance(m m ol/m 2 s) a b c Table 1. Results from paired t-test (H 0 : µ D =0, α=0.05) showing the agreement between the natural log transformed stomatal conductance data obtained by the LI-1600, the SC-1, and the AP-4. Data were obtained in field conditions with paired samples having identical environmental conditions. Statistics on the non log transformed data are also included. Fig. 1. Correlation plots containing one-to-one lines of the regression of the natural log of the stomatal conductance data obtained from each instrument are shown on the left while corresponding plots showing matched pair stomatal conductance means and matched pair stomatal conductance differences for each instrument are shown on the right. Plots a and d compare the LI- 1600 and the SC-1, plots b and e compare the LI- 1600 with the AP-4, and plots c and f compare the AP-4 with the SC-1. Results Figure 1 shows the agreement of stomatal conductance values obtained from the AP-4, the LI- 1600, and the SC-1 as well as the differences between each mean pair. After normalizing all of the data using log transformation, a regression of SC-1 verses the AP-4 (Fig 1-c) reveals that the SC-1 generally measures slightly lower conductance readings than the AP-4 (slope of 0.9701 and an R 2 of 0.62 (n=235). A regression of the formerly mentioned instruments against the LI-1600 suggests that both the AP-4 (Fig 1-b) and the SC- 1 (Fig 1-a) give conductance readings above that of the LI-1600 with slopes of 1.0202 and 1.1182, respectively and R 2 of 0.45 (n=155) and 0.52 (n=129), respectively. Mean difference graphs (Figure 1 d - f) show the trends in instrument differences from low to high conductivity values. These data suggest that high conductance values may incorporate more measurement error that low conductance values. However, not enough high conductance values are available to make any specific conclusions. Methods (contd.) We tested our first assumption by measuring spatial variability of stomatal conductance on a leaf using all three instruments. For large leaves, stomatal conductance was measured at six points on the leaf surface. The second assumption is certainly not completely valid. Not only do the effects of previous porometer measurements likely affect stomatal conductance, the environmental conditions controlling stomatal conductance often change minute-by-minute. Stomatal conductance data were normalized using natural-log transformation. A paired t-test was used to determine the confidence intervals of the difference between instruments. Results (contd.) Results (contd.) Conclusions 1) All instruments show significant systematic differences in measured stomatal conductance at the α=0.05 level. However, these differences do not invalidate comparisons between conductance measurements between the three porometers. 2) All three instruments appear to have poorer agreement at high conductance values as shown in plots d, e, and f. However, there are not enough data at high conductance values to statistically validate these conclusions. 3) The LI-1600 generally reads lower than the other two porometers; especially at high conductivity values. Similar results have been found in other research (Delta T Devices, 2006). 4) The AP-4 and the SC-1 agree better than either of the two agree with the LI- 1600. 5) Considerable spatial variability in conductance over the surface of single leaf and changing environmental conditions likely cause scatter in the intercomparison data. These errors can be minimized in practical application through the use of replicated measurements. 6) Results suggest that measurements should be comparable among the AP-4 and SC-1 at all conductance levels. The LI-1600 tends to measure lower conductances than the other two porometers, especially at higher conductances. This should be corrected for if results obtained by the two Reference d -1500 -1000 -500 0 500 1000 1500 0 500 1000 1500 2000 M atch PairM ean Conductance m mol/m2s(LI-1600,SC-1) M atched PairDifference(LI-1600 -SC-1) -1500 -1000 -500 0 500 1000 1500 0 500 1000 1500 2000 M atch PairM ean Conductance m m ol/m 2s(LI-1600, A P-4) M atched PairDifference(LI-1600 -AP-4) e d -1500 -1000 -500 0 500 1000 1500 0 500 1000 1500 2000 M atch PairM ean Conductance m m ol/m 2s(A P-4, SC-1) M atched PairDifference(AP-4 -SC-1) f Delta T devices. Draft Leaf Porometer (SC- 1) Evaluation. Unpublished manuscript. October 2006. Instrum ents Sam ple size (N ) A verage Difference Standard Deviation from the M ean Standard Error from the M ean 95% C onfidence Interval P-value of T-test LI-1600 -SC-1 129 -0.61 0.52 0.05 (-0.70,-0.52) <0.005 LI-1600 -A P-4 155 -0.62 0.65 0.05 (-0.72,0.52) <0.005 A P-4 -SC-1 198 0.14 0.51 0.04 (0.07, 0.02) <0.005 LI-1600 -SC-1* 129 -192 235 20.7 LI-1600 -A P-4* 155 -241 350 28.1 A P-4 -SC-1* 198 79 279 19.8 * Values are descriptive statistics only. Stomatal conductance measurements at different locations on single, fully sunlit leaves show that significant spatial variability is present over the leaf surface (Figure 2). Doug Cobos, Decagon Devices (509) 332-2756 [email protected] Methods

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Page 1: Introduction Stomatal conductance regulates the rates of several necessary processes in plants including transpiration, carbon dioxide assimilation, and

Introduction

Stomatal conductance regulates the rates of several necessary processes in plants including transpiration, carbon dioxide assimilation, and respiration. Both steady state and dynamic porometers are currently used to measure stomatal conductance. However, few tests or measurements have been performed to compare the results obtained with the various porometers. It is important for scientists to understand the performance of each porometer in order to compare stomatal conductance data gathered from various models. In this study, we focused on three commercially available porometers: the AP-4 (Delta-T Devices, Cambridge, UK), the SC-1 (Decagon Devices, Pullman, WA), and the LI-1600 (Li-Cor, Lincoln, NE). The objective of this study was to understand the relationship between the stomatal conductance data gathered with the different instruments

An Inter-comparison of Three Commercial An Inter-comparison of Three Commercial PorometersPorometers

L. Bissey1, D. Cobos2, C. Campbell2

1. School of Earth and Environmental Science, Washington State University, 2. Decagon Devices, Inc.

Figure 2. Example data showing leaf-level stomatal conductance variability. Numbers in black, blue, and purple are conductance data obtained using the AP-4, SC-1, and LI-1600 respectively. Data from different instruments were collected at different times and should not be compared among porometers.

962, 673, 410

678, 674, 362

701, 560, 383

690, 918, 425

753, 573, 322

1053, 964, 356

We measured stomatal conductance of approximately 200 leaves of various species with an AP-4, SC-1, and LI-1600 under field conditions. Changes in environmental conditions were minimized by taking measurements with the three porometers on similar locations of each leaf in rapid succession. On each leaf, stomatal conductance measurements with the three porometers were conducted in random order to prevent systematic bias from previous porometer measurements. Each porometer was factory calibrated within 60 days of the beginning of measurements.

This methodology assumes that: 1) stomatal conductance was measured on the exact same footprint on each leaf, and 2) environmental factors (radiation, temperature) didn’t change over the 1-2 minute timescale needed to make the three measurements on each leaf.

1

2

3

4

5

6

7

8

1 2 3 4 5 6 7 8

Ln LI-1600 Stomatal Conductance (mmol/m2s)

SC-1

Sto

mat

al C

ondu

ctan

ce (m

mol

/m2 s)

1

2

3

4

5

6

7

8

1 2 3 4 5 6 7 8

Ln LI-1600 Stomatal Conductance (mmol/m2s)

LN A

P-4

Stom

atal

Con

duct

ance

(mm

ol/m

2 s)

1

2

3

4

5

6

7

8

1 2 3 4 5 6 7 8

LN AP-4 Stomatal Conductance (mmol/m2s)

LN S

C-1

Sto

mat

al C

ondu

ctan

ce (m

mol

/m2 s)

a

b

c

Table 1. Results from paired t-test (H0: µD=0, α=0.05) showing the agreement between the natural log transformed stomatal conductance data obtained by the LI-1600, the SC-1, and the AP-4. Data were obtained in field conditions with paired samples having identical environmental conditions. Statistics on the non log transformed data are also included.

Fig. 1. Correlation plots containing one-to-one lines of the regression of the natural log of the stomatal conductance data obtained from each instrument are shown on the left while corresponding plots showing matched pair stomatal conductance means and matched pair stomatal conductance differences for each instrument are shown on the right. Plots a and d compare the LI-1600 and the SC-1, plots b and e compare the LI-1600 with the AP-4, and plots c and f compare the AP-4 with the SC-1.

Results

Figure 1 shows the agreement of stomatal conductance values obtained from the AP-4, the LI-1600, and the SC-1 as well as the differences between each mean pair. After normalizing all of the data using log transformation, a regression of SC-1 verses the AP-4 (Fig 1-c) reveals that the SC-1 generally measures slightly lower conductance readings than the AP-4 (slope of 0.9701 and an R2 of 0.62 (n=235). A regression of the formerly mentioned instruments against the LI-1600 suggests that both the AP-4 (Fig 1-b) and the SC-1 (Fig 1-a) give conductance readings above that of the LI-1600 with slopes of 1.0202 and 1.1182, respectively and R2 of 0.45 (n=155) and 0.52 (n=129), respectively. Mean difference graphs (Figure 1 d - f) show the trends in instrument differences from low to high conductivity values. These data suggest that high conductance values may incorporate more measurement error that low conductance values. However, not enough high conductance values are available to make any specific conclusions.

Methods (contd.)

We tested our first assumption by measuring spatial variability of stomatal conductance on a leaf using all three instruments. For large leaves, stomatal conductance was measured at six points on the leaf surface.

The second assumption is certainly not completely valid. Not only do the effects of previous porometer measurements likely affect stomatal conductance, the environmental conditions controlling stomatal conductance often change minute-by-minute.

Stomatal conductance data were normalized using natural-log transformation. A paired t-test was used to determine the confidence intervals of the difference between instruments.

Results (contd.) Results (contd.)

Conclusions1)All instruments show significant

systematic differences in measured stomatal conductance at the α=0.05 level. However, these differences do not invalidate comparisons between conductance measurements between the three porometers.

2)All three instruments appear to have poorer agreement at high conductance values as shown in plots d, e, and f. However, there are not enough data at high conductance values to statistically validate these conclusions.

3)The LI-1600 generally reads lower than the other two porometers; especially at high conductivity values. Similar results have been found in other research (Delta T Devices, 2006).

4)The AP-4 and the SC-1 agree better than either of the two agree with the LI-1600.

5)Considerable spatial variability in conductance over the surface of single leaf and changing environmental conditions likely cause scatter in the intercomparison data. These errors can be minimized in practical application through the use of replicated measurements.

6)Results suggest that measurements should be comparable among the AP-4 and SC-1 at all conductance levels. The LI-1600 tends to measure lower conductances than the other two porometers, especially at higher conductances. This should be corrected for if results obtained by the two instruments are to be compared.

Reference

d

-1500

-1000

-500

0

500

1000

1500

0 500 1000 1500 2000

Match Pair Mean Conductance mmol/ m2s (LI-1600, SC-1)

Mat

ched

Pai

r Dif

fere

nce

(LI-

1600

- SC

-1)

-1500

-1000

-500

0

500

1000

1500

0 500 1000 1500 2000

Match Pair Mean Conductance mmol/ m2s (LI-1600, AP-4)

Mat

ched

Pai

r Dif

fere

nce

(LI-

1600

- A

P-4)

e

d

-1500

-1000

-500

0

500

1000

1500

0 500 1000 1500 2000

Match Pair Mean Conductance mmol/ m2s (AP-4, SC-1)

Mat

ched

Pai

r Dif

fere

nce(

AP-

4 - S

C-1

)

f

Delta T devices. Draft Leaf Porometer (SC-1) Evaluation. Unpublished manuscript. October 2006.

InstrumentsSample size (N)

Average Difference

Standard Deviation from the

Mean

Standard Error

from the Mean

95% Confidence

Interval

P-value of T-test

LI-1600 - SC-1 129 -0.61 0.52 0.05 (-0.70, -0.52) <0.005

LI-1600 - AP-4 155 -0.62 0.65 0.05 (-0.72, 0.52) <0.005

AP-4 - SC-1 198 0.14 0.51 0.04 (0.07, 0.02) <0.005

LI-1600 - SC-1* 129 -192 235 20.7

LI-1600 - AP-4* 155 -241 350 28.1

AP-4 - SC-1* 198 79 279 19.8

* Values are descriptive statistics only.

Stomatal conductance measurements at different locations on single, fully sunlit leaves show that significant spatial variability is present over the leaf surface (Figure 2).

Doug Cobos, Decagon Devices(509) [email protected]

Methods