Agricultural and Forest Meteorology, 54 ( 1991 ) 241-261

241

Elsevier Science Publishers B.V., Amsterdam

Estimation of maize (Zea m a y s L. ) canopy

conductance by scaling up leaf stomatal
conductance*
P. Rochette a, E. Pattey b, R.L. Desjardins a, L.M. Dwye#, D.W. Stewart a and
P.A. Dub6 b
aLand Resource Research Centre, Research Branch, Agriculture Canada, Ottawa, Ont. KIA 0C6,
Canada
bD~partement de Phytologie, Facult~ des Sciences de l'Agriculture et de l'Alimentation, Universitk
Laval, Quebec City, Que. GIK 7P4, Canada
(Received 1 November 1989; revision accepted 22 October 1990)

ABSTRACT
Rochette, P., Pattey, E., Desjardins, R.L., Dwyer, L.M., Stewart, D.W. and Dubr, P.A., 1991. Estimation of maize (Zea mays L.) canopy conductance by scaling up leaf stomatal conductance. Agric.
For. Meteorol., 54: 241-261.
Transpiration is partially controlled by the plant at leaf level through the degree of aperture of the
stomata. Mathematical models estimating the transpiration of plant stands using a conductance network approach to water vapor transfer thus need a plant surface control term (g~). Techniques involving different degrees of simplification of canopy structure have been proposed to estimate gs from
measurements or estimates of leaf stomatal conductance (gs). This study compares the performance
of some of these techniques by examining the patterns of diurnal and seasonal variation for a maize
crop grown at Ottawa, Canada. Measurements ofg~ were made for sunlit and shaded leaves at three
levels in the plant canopy and the response of gs to photosynthetic photon flux density (Qp) has been
parameterized. Values ofg~ obtained by different scaling-up methods were compared among themselves and with those obtained from the Penman-Monteith equation (g).
The main results and conclusions were: daily maximum gs values of sunlit leaves generally occurred
on or slightly before maximum radiation; the response of gs to Qp was a function of leaf levels in the
canopy and growth stage; the response ofgs to low Qp increased with leaf area index for shaded leaves;
modelling of gs based on Qp, leaf temperature and leaf water potential failed to give good estimates
under overcast afternoon conditions; measurement of g~ on horizontal portions of leaves led to an
overestimation of g~; spherical leaf angle distribution assumption gave the best estimates of g~; the
shelter factor (ratio of scaled-up g~ over gc ) tended to increase as the ratio of the canopy aerodynamic
conductance to top leaf stomatal conductance increased.

INTRODUCTION

Energy and mass fluxes to and from a plant canopy are the result o f turbulent transport processes that are highly variable in time and space. In the last
*LRRC Contribution No. 89-72.

0168-1923/91/$03.50

1991 - - Elsevier Science Publishers B.V.

242

P. ROCHETTE ET AL.

ten years, it has been shown that the traditional treatment of these fluxes by
the flux-gradient theory has little in common with the real physical processes
involved (Finnigan, 1979; Denmead, 1984; Finnigan and Raupach, 1987).
More realistic approaches using higher order closure and Lagrangian models
are being investigated, but their use requires the parameterization of important terms and they are not yet usable on a practical basis (Finnigan and Raupach, 1987 ). Therefore, many operational models express the effect of molecular and turbulent diffusion on the magnitude of exchanges by alternative
approaches, such as an analogy with electrical conductance.
These models describe the path of water vapor from the soil and leaf surface to the atmosphere above the canopy through a network of conductances.
Many of them estimate evapotranspiration of a plant stand through an approach involving the leaf energy budget in which the canopy is considered to
act as 'big leaf'. Despite the limitations of this approach (Philip, 1966; Denmead, 1984), Jarvis et al. (1981) concluded that assumptions inherent in
reducing the plant canopy to two horizontal dimensions induce errors in transpiration estimates of only a few percent.
Models based on the energy budget using the 'big-leaf' assumption include
many meteorological and canopy variables. The meteorological variables are
readily provided by appropriate instrumentation, but the canopy variables
are more difficult to obtain. Among them, the determination of stomatal conductance (gs) is most difficult. Problems in the measurement of gs, and the
requirement for the simulation of mass and energy exchanges of terrestrial
ecosystems, have resulted in the development of sub-models for the estimation of g~. The stomatal response to environmental variables has been investigated (Jarvis, 1976) and some specific relationships have been suggested
for maize (Stewart, 1970; Beadle et al., 1973; Shawcroft et al., 1974; Turner,
1974; Norman, 1979). However, no mechanistic model has yet been proposed that considers all environmental variables and their interaction (Jarvis
et al., 1981 ), and Squire and Black ( 1981 ) stressed that models must be calibrated for the age and general conditions of the crop under study.
The energy combination approach to evaporation from canopies includes
a canopy conductance (gc). Jarvis et al. (1981) proposed an estimation of
the canopy conductance, defined as the sum of the g~ values of all individual
leaves in an imaginary column through the canopy standing on a unit ground
area (g~). Different sampling procedures involving the stratification of samples, together with proper weighting according to the corresponding fractional leaf area index (L), have therefore been used (Leverenz et al., 1982;
Whitehead et al., 1984; Beadle et al., 1985 ).
The objectives of this paper are: ( 1 ) to quantify the magnitude of the difference between g~ (obtained by different scaling-up methods) and gc as calculated from the Penman-Monteith equation; (2) to propose a shelter factor
to predict gc from gs- This study also identified diurnal and seasonal variation

ESTIMATION OF MAIZE CANOPY CONDUCTANCE

243

of maize gs as a function of leaf level and solar exposure (sun or shade ), and
parameterization of the g~ response to Qp for different leaf levels and growth
stages.
MATERIALS AND METHODS

Data collection
Canopy parameters
Measurements of stomatal conductance were made on an hourly basis in a
40 ha maize field during 7 days distributed over the 1988 growing season at
Ottawa, Canada. Measurements were made on sunlit and shaded leaves at
three levels in the canopy. Five observations were made for each solar exposure, level and hour. The levels were defined in such a way that they represented the lower 25%, the middle 50% and the top 25% of the canopy L. The
same portable photosynthesis system (LCA2, Analytical Development Co.
Ltd., Hoddesdon, U K ) was used on all days except 18 August, when measurements were taken with a LI-COR 6200 (LI-COR Inc., Lincoln, NE). Values ofgs were calculated assuming equal numbers of stomata on both sides of
the leaf.
Pertinent canopy measurements (plant height, leaf position and area) were
made on a regular basis throughout the growing season. The water potential
of leaves in the sun and in the shade was measured with a pressure bomb, at
all three levels on three of the seven measurement days. Three measurements
were made for each solar exposure and level during each hour.
All measurement days were cloudless except 18 July, which had uniform
cloud cover beginning around midday after a cloudless morning.

Meteorological parameters
The following parameters were recorded every 10 s and averaged on an
hourly basis: net radiation (Rn) with two net radiometers (CN 1, Middleton
Instruments, Vic., Australia), direct and diffuse solar radiation (Rs) and Qp
with photoelectric sensors (LI-200SA and LI-190SA, LI-COR Inc., Lincoln,
NE ) and shadow bands, soil heat flux (G) with four plates (CN3, Middleton
Instruments, Vic., Australia), wind velocity and air temperature (Ta) with a
sonic a n e m o m e t e r - t h e r m o m e t e r (DAT-310, Kaijo Denki Ltd., Tokyo), and
air temperature and humidity (Probe 207, Campbell Scientific, Logan, U T ) .
Average values were calculated when more than one sensor was used for the
same parameter. Sensible heat, latent heat and m o m e n t u m fluxes were measured over the same interval by the eddy correlation technique in which the
water vapor fluctuations were monitored by an open-path CO2 and H20 fastresponse sensor (Chahuneau et al., 1989; Desjardins, 1990).
Shaded and sunlit fractions of L, and direct and diffuse Qp at different lev-

244

P. ROCHETTEET AL.

els in the canopy for a horizontal plane and for five leaf inclination angle
classes were calculated from radiation above the canopy, solar elevation and
canopy characteristics using CUPRAD, the radiation subroutine of the
CUPID model (Norman, 1979). A spherical foliage element distribution, a
leaf grouping factor of 0.85 and an L fraction per canopy layer of 0.2 were
used in CUPRAD.

Analysis
Reference surface conductance (gc)
The reference surface conductance to water vapor (gc) was obtained by
rearranging the Penman-Monteith equation (Monteith, 1965 )
JTl'E~ga'H
gc - - S ( g n - G) "l-paCp(e* (Ta) - ea)ga,H --/LE(s q- 7)

where
ga,H =

1 +

ga,M

,)'

(])

(2)

gb,H

ga,M --paU(Z)

(3)

ku*
gb'n --ln (Zo/Zn)
gb,H--

(4)
for ZH = 0.2Z0

where 2 is the latent heat of vaporization for water vapor, Eis the evaporation
flux, ~, is the psychrometric constant, g~,U and g~,M are the aerodynamic conductances for sensible heat and momentum, respectively, s is Oe*/OTa, Pa is
the air density, cp is the specific heat of air at constant pressure, e* (Ta) is the
saturation water vapor pressure at T~, ea is the actual water vapor pressure,
gb,u is an additional conductance for water vapor to account for form drag at
the surface (Thom, 1975), z is the flux of momentum, u(z) is the wind velocity at height of reference, u* is the friction velocity, k is von Karman's
constant (0.41) and Zo and Zn are the roughness lengths for momentum and
sensible heat, respectively.

Parameterization of gs
Jarvis (1976) proposed that gs could be estimated using the maximum gs
(gs,x) for the plant species under study, corrected for non-optimum leaftem-

ESTIMATIONOF MAIZECANOPYCONDUCTANCE

245

perature (T1), leaf water potential (tl) and vapor pressure deficit (D) by the
following approach
gs=gs,xf~ (~l) f z ( T1) f 3 ( O )

(5)

where no interaction is assumed between the parameters.

In this experiment, gs,x was estimated by the stomatal response to Qp in the
early part of the day (before 12:00 EST) when plants were not exposed to
moistur~ or heat stresses. The response ofgs to Qp in field conditions has been
described by a rectangular hyperbola following the approach proposed by
Dwyer and Stewart ( 1986 ) for photosynthesis
a Qp
g~ - 1 + Qp [ o~/ ( g~2ooo - ge ) - 1/2000] t-g~

(6)

where a is the initial slope of the curve (at Qp= 0), g~ is the maize leaf cuticular conductance (0.333 m m s-~ ) and gs2ooois the stomatal conductance at
a Qp of 2000/~mol m - 2 s- ~.
Norman (1979) used the following correction functions for corn
A (~t,) = (, - ~1,2)/ (~ffl,i -L2) for ~t,.i > , >,,2

(7)

where ~,~is the threshold at which stomata begin to close and l,Z is the value
at which all stomata are closed. The function is set to unity for ~'l values greater
than ~UL1and to zero for 1 values smaller than ~'1.2. Values of - 10 and - 2 2
bars were chosen for el,1 and L2, respectively (Sellers and Dorman, 1987).
1
f2(rl)={l'lt-~
(-rl'--Tl')16m~ - forT,>T,o
(8)
m( T,,u - Tl,o)_] )
6
--1
, F (Z,o-r,) l
m~ for rl <TLo
(8)
where Tl,o is the optimum leaf temperature corresponding with the maximum
conductance, Tl,u and TI,Lare, respectively, the upper and lower leaf temperature limits above and below which there is no stomatal opening, and m is a
constant. Values of 15, 30 and 45 C were chosen for T~,L,Tl,o and TLu, respectively, and m was set equal to 5 (Norman, 1979 ). No correction function was
used to take into account the effect of D on g~.
Scaling-up techniques
Six different methods to scale up from leaf stomatal conductance values to
canopy conductance have been tested.
(i) Bulk average (BA) (Bailey and Davies, 1981 )
gs=[(gs,.+gs,e)/2]xL

246

P. ROCHETTE ET AL.

where &,. and &,e are, respectively, the mean & of sunlit and shaded horizontal leaves over the whole canopy.
(ii) Top sunlit layer sampling (TS) (Whitehead et al., 1981 )
gs =gs,n,v L
where g~,,,a-is the mean g~ of the sunlit horizontal leaves in the top 25% of the
L.
(iii) Weighted type (W) (Duncan et al., 1967; Sinclair et al., 1976)
gs = g s , n

L . +gs,e Le

where L. and L e are, respectively, the total sunlit and shaded L over the whole
canopy.
(iv) Effective leaf area index (EL) (Szeicz and Long, 1969 )
& = [ (&,, +gs,e)/21XLf
where the effective L (Lf) is

Lf=L

ifL<-Lx/2

Lf = Lx/ 2 if L > Lx/ 2

where Lx is the maximum L expected for the canopy under the conditions of
the study.
(v) Horizontal canopy layer (HL)
3
gs = E
i=1

(Ln,iXgs,ni+Le,iNgs,e,i)

where i represents the canopy layers.

(vi) Multiple leaf angle classes canopy layer (MAL) (Sellers et al., 1986 )
gs = ~

i=l

=1

(g~,~,ijxL~,i,j)+gs,e,~XLe,i

where i represents the canopy layers, j is the leaf inclination angle classes and
gs values are estimated using eqn. ( 5 ). All other methods used g~ values measured on horizontal sections of leaves.
RESULTS A N D D I S C U S S I O N

Diurnal and seasonal variation of stomatal conductance (gA

Diurnal variations in gs of shaded and sunlit leaves at three levels in the
maize canopy are presented in Fig. 1 (a) and 1 (b) for the seven measurement
days. Typical mid-day values for the standard error of the mean were < 15%
of the mean.

ESTIMATION OF MAIZE CANOPY CONDUCTANCE

247

Shaded leaves generally had gs values less than half those of sunlit leaves at
the same level, except on 8 and 29 July. The greater difficulty in sampling the
shaded leaves in sparser canopies probably caused the high middle and top
leaf values on 8 July. Shade periods are shorter in such canopies and shaded
leaves may not be in equilibrium with their light environment at sampling
time. Relatively high gs on shaded leaves at the top of the canopy on 29 July
seems to have been caused by a higher gs response to low radiation intensity
(or parameter in Table 1 ) at this development stage (silking). Measurements
on shaded leaves at the top of the canopy taken on 27 July (data not shown)
were also higher than those taken earlier or later in the season.
Diurnal variation in gs of shaded leaves usually followed a pattern of a maxi m u m early in the day and a gradual decrease toward the end of the day. Early
measurements on 18 July and 8 August were deleted because dew on the leaf
surfaces resulted in erroneously high values.
The seasonal trend in g~ of shaded leaves is a decrease from the beginning
to the end of the season. This tendency is probably the result of a decrease in
the diffuse radiation intensity as the canopy gradually becomes more dense,
followed by leaf senescence later in the season.
Values of g~ of sunlit leaves were generally smaller at the bottom of the
canopy, while values were approximatively the same for the middle and top
of the canopy. An exception occurred on 21 June when similar values were
found at all levels in the canopy. On that date, the bottom leaves were as
active as the higher leaves (similar level of photosynthesis) and the incomplete canopy did not restrict radiation to the lower level. Later in the season,
both the decline in radiation and the earlier aging of lower leaves explain the
smaller values at the bottom of the canopy.
Diurnal variation in g~ was characterized by a m a x i m u m that occurred later
in the day for sunlit leaves than for shaded leaves. M a x i m u m g~ generally
corresponded with the time of m a x i m u m radiation or occurred slightly before
it (Fig. 2 ). Morning values ofg~ were usually higher than afternoon values for
a similar Qp. This indicates that other limiting factors played a significant
role. Diurnal variation in gs of sunlit leaves was typical of plants grown in
h u m i d environments. They did not show the consistent mid-day decrease reported in drier environments (Tenhunen et al., 1987). However, weak midday stomatal closure seems to have occurred on 8 July. The highest air temperatures and vapor pressure deficits recorded during the study occurred on
this day (Fig. 2 ).
No consistent seasonal pattern was observed for sunlit leaves. This may be
because sunlit leaves are more vulnerable to water stress than shaded leaves
and such temporary stress conditions may mask the effect of longer term factors such as leaf aging.

248

P. ROCHETTE ET AL.

Parameterization Ofgs
Table 1 lists the values of the coefficients required to estimate gs,x (eqn.
( 5 ) ) based on morning data for each level and for the entire canopy for 6
days of the experiment. Curves for 29 July are shown as examples of the fitting procedure (Fig. 3 ).
Variations of gs2ooo from day to day followed those of the maximum measured values of g~ on sunlit leaves at each level, while canopy coefficients
averaged out the differences between levels (Table 1 ). Field ( 1987 ) reported
that gs of Lepechinia calycina at high Qp peaked in mid-season and that g~
response to low Qp was nearly constant throughout the growing season, with

(a)

BOTTOM PART OF CANOPY

20

- - - - 21/06
08/07
- - - 13/07
. . . . 18/07

16
12

20

SHADE

SUN :16

i-~

/\

12

\j\

20

11 13 15 17
9 11 13 15 17 1
MIDDLE PART OF CANOPY
SHADE

16

20

SUN

12

12

gffl8

8
4

4
7

........
9

20

..........
? 0
11 13 15 17 7 9 11 13 15 17 19
TOP PART OF CANOPY
20
SHADE[

16

SUN :16

12 ~
8:

11 13 15 17 7 9
t (h)

11 13 15 17 19

Fig. 1. (a) Stomatal conductance (gs) at three levels in a maize canopy during 4 days in the
1988 growing season (Ottawa, Canada). (b) Stomatal conductance (gs) at three levels in a
maize canopy during 3 days in the 1988 growing season (Ottawa, Canada).

ESTIMATION

OF

MAIZE

CANOPY

b)

BO3-FOM PART OF CANOPY

20
---....

16

249

CONDUCTANCE

29/07 SHADE
08/08
18/08

12

SUN

16

/" \

i'
,

20

12

,\

"--

~ "fl" f3' 1"5" 1"7" t

MIDDLE PART OF CANOPY

20.

20

SHADE

16

SUN

/ , ,L

'~ 12

1"1 " 1"3 f5 1"719

16

12

8:

41

oi

" 1"1 1"3" 1"5" 1"7" 7 " ~ " 1"1 " 1"3" 1"5" 1 " 7 " 1 9

TOP PART OF CANOPY

20

20

SHADE

16

SUN

16

f- ..~P\

12

.1"-~ / "k.~
,,,,,

/\

12

",%

11 13 15 17

11 13 15 17

t (h)

the exception of a period shortly after leaf emergence ( 14 days) when lower
values occurred. No consistent seasonal trend in gs2ooowas found in this study.
However, the values of a increased gradually during July and returned to
early season levels on 18 August This suggests that the response ofgs of shaded
leaves to low Qp increased as the plant canopy became denser, but that the
process was reversed in August because of leaf aging.
Reported m a x i m u m conductances of maize g~ in response to Qp are approximately 10.0 m m s- ~ (including both leaf faces) (Turner and Begg, 1972;
Uchijima, 1976; N o r m a n and Campbell, 1983 ). Estimates ofgs2ooo integrated

250

P. ROCHETTE ET AL.

(a)
40
35

30

\ JUNE 21

1~

25 y
20 ' ,
15

'

\
'

.-I

""

~
\

2000

40
35

1600

30 /

1200
800

- /- /-~- - ' 8~ - - - - - - , ~ J U L Y

2000

, .~-..

25'

20.

,,'

15.

1600

'

1200

\
\

800
\

10
5

- - - - Qp
Ta

\
\.

400

10, 7"
5.

400

04

o
~"
o

..........

~
~

==

11 13 15 17 19

...........
7

E
oE
~.

11 13 15 17 19

40.

,0

E3 35

30

-~. JULY 13

2000

35' JULY 18
/~

2000

1600

30"

1600

25 ~
/

20
15

~," "

1200
\

10
,
5 "
7

~k'--~.~

25'
800

15. I

400

10.
5

,,

...........

\
\~

...........
0
9 11 13 15 17 19

t (h)

1200

20'

, A" " . .
,"

600
"-

~,~

400
0

11 13 15 17 19

t (h)

Fig. 2. (a) Diurnal pattern of photosynthetic photon flux density (Qp), air temperature ( T a)
and vapor pressure deficit (D) during 4 days in the 1988 growing season (Ottawa, Canada).
(b) Diurnal pattern of photosynthetic photon flux density (Qp), air temperature (Ta) and vapor pressure deficit (D) during 3 days in the 1988 growing season (Ottawa, Canada).

over all canopy levels varied between 7.2 and 13.0 m m s- ~ in this study (Table 1 ). It can be seen that the utilization of a single set of coefficients throughout the season could result in a significant error. Special attention should,
therefore, be attached to the parameterization of gs with respect to leaf aging
and leaf history when modeling gs in the field.
Values of gs2oooincreased 20-100% from the bottom to the top of the canopy. Smaller g~ values in the lower part of the maize canopy agree with earlier
studies, but those studies found larger vertical gradients in the upper half of
the canopy (Turner, 1974; Uchijima, 1976).

Canopy conductance (gA

Surface conductance (go) was estimated by the P e n m a n - M o n t e i t h equation (eqn. ( 1 ) ) and is summarized for the seven days (Fig. 4). It was as-

ESTIMATIONOF MAIZECANOPYCONDUCTANCE

251

4O
35l JULY29

(b)

12000

; ~ : : ] / ~

~,,ll

-"~" t.oo |

o 1o
o,

t1200

\\\

....

X",

,oo

. . . . . . . . . . .

,o

11 13 15 17 19

t (h)
40"
2000 35'

::I ,,ous,.

leoo 3oi

.-

1600 ,-,

'7,
m

1200

20'

1200 ~E

8OO

15.

8OO

4O0

9 11 13 15 17 19
t (h)

25'

,-, lO 1

2000

AUGUST 18

10'
5'
0

E
O.

ji

~*

4OO 0

11 13 15 17 19
t (h)

TABLE 1
Leaf area index (L), slope of the relationship between stomatal conductance (gs) and photosynthetic
photon flux density ((29) at (2v = 0 (or), and value of gs at (2p = 2000/tmol m - 2 s - ~ ( gs2ooo)
Day

21 June
8July
13 July
18 July
29 July
8 August
18 August

0.4
1.8
2.5
2.6
2.7
2.7
2.7

O/ (X10 -2) (mm m2/tmol -~ )

Whole
canopy

Bottom

.
.
2.28
4.96
6.04
10.54
8.69
2.56

.
2.98
4.93
7.84
8.60
7.06
3.17

aApproximate value.

g,2ooo ( m m s -~ )

Middle

Top

2.12
4.65
7.42
10.64
9.20
2.23

2.11
3.62
3.99
11.31
6.73
2.77

Whole
canopy

Bottom

Middle

Top

10.0 a
12.3
7.4
8.7
9.8
10.9
7.2

10.0 a
8.1
6.0
6.2
5.8
9.8
5.5

10.0 a
12.9
8.9
8.8
9.2
11.6
8.0

9.5 a
13.5
7.2
10.8
12.5
11.3
8.0

252

P. ROCHETTEET AL.
24
22
20
18
16
14
12
10
8
6
4
2

24.
2 2 BOTTOM OF CANOPY
20
18
16
14
12
10

'
4

i-"

||

r'L

E
E

12

MIDDLE OF CANOPY

|
I

I
|
II

r = .50

16 20

24

II

12

16

20

24

2,~

22 ]
2O J
184
164
144 ~
124"
104 JU.|

22
TOP OF CANOPY
20
18
Ii
16
14
12'
10.
8
trKV
6'
4
2'
r = .57
0

12

ALL LEVELS

'

,'

16 20

,
~

',-.,'..:

84,,
44

,,,

"
i

,,,.

zip

2|
r = .56
01 . . . . . . . . . . .
24
0
4
8 12 16 20

24

Q p ( p m o l m'2s'Ix 10"2)

Fig. 3. Stomatal conductance (gs) response to photosynthetic photon flux density (Qp) at three
levels in a maize canopy on 29 July 1988 at Ottawa.
50
21106
08O7
13/07
18/07
29/07
08/08
18/08

45
- - . ....
---

40.
35 . . . . .

----

zo

E 2s.
"-'20-

"'./.
~ I
./"
l.J-

~)

o115,

/ ~" \

r...~.~l
"///\

-.

"

\
\

10.
5O,

'

1'1

1'3 ' 1'5 ' 1'7 ' 19

t (h)

Fig. 4. Surface conductance to water vapor transfer (g) obtained from the Penman-Monteith
equation in a maize field in 1988 (Ottawa, Canada).

ESTIMATION OF MAIZE CANOPY CONDUCTANCE

2 53

sumed that evaporation from the soil surface was insignificant. This condition was met on all days except 29 July, when direct evaporation from the soil
surface could not be ignored because of rain on the preceeding days. Values
of g~ peeked in mid-season with lower values in early July and mid-August.
Low 21 June values were caused by the large contribution of a dry soil surface
since L was low (Table 1 ) and little rain was received in the previous weeks.

Scaling up of gs
Values ofgs were scaled up using the six methods, described earlier, to provide gs. Figure 5 (a) and 5 (b) compares g~ values estimated using the six
methods to the calculated reference values (gc). The BA, W and HL methods
gave very similar estimates for every day of the experiment. This suggests that
not distinguishing between sunlit and shaded L in the weighting procedure
does not produce different gs values for m e d i u m density canopies, like the
one studied, if sampling is made in the sun and in the shade at different levels
in the canopy. This conclusion could be different for a denser or a sparser
canopy, where sunlit and shaded L fractions over the whole canopy are significantly different, and when sampling is random or incomplete.
Sampling only the top sunlit leaves (TS) led to gs values that were up to
double those of the three methods discussed above. The use of effective L
(EL) estimated g~ values that were smaller than those given by the other scaling-up methods. This difference became constant as the L reached its maxim u m value.
Canopy conductance estimates obtained by the BA, W and HL methods
were larger than gc on 8 and 13 July, and 18 August. The EL method gave g~
values close to gc on 8 and 13 July and on 8 August, and smaller values on the
other days. The TS m e t h o d produced g~ values that exceeded gc on all days
except 29 July. It, therefore, seems that scaling-up methods that require a
weighting based on sunlit and shaded L and canopy level tend to overestimate
g~ in this study. The major reason for this is that field measurements were
made on horizontal sections of leaves. This procedure assumes that horizontal leaves are exposed to a light regime close to that of the average leaf. However, for a spherical leaf distribution, this would lead to a systematic overestimation, especially in the midday period.
On 21 June, the BA m e t h o d was computed assuming that all the leaves were
sunlit. The omission of the shaded leaves also contributed to the overestimation of g~ and is partly responsible for the fact that the scaled-up g~ values by
the BA method were higher than g~ values.
Values ofg~ obtained by the EL m e t h o d had a good correspondence with gc
values when the canopy was incomplete, but the method fails to estimate gc
when the canopy is fully developed. However, early in the season and for dry
soil surfaces, the EL method gave the best approximation of go. The TS method
consistently overestimated gc, as was suspected by Whitehead et al. ( 1981 ).

254

P. ROCHETTEET AL.

However, under wet soil surface conditions, TS gave the best estimates of gs
(29 July).
Values of gs obtained with the MAL method were systematically smaller
than those produced from the BA, W and HL methods. The difference was
larger early ( 8 and 13 July) and late ( 18 August) in the season. This method
produces gs values that were closer to gc early and late in the season. The MAL
method weights g~ values according to a spherical distribution of foliage elements in the canopy. The improvement ofgs obtained by MAL early and late
(a)
40

40

BA
J U N E 21

- - - TS

30 i . . . . . W
- - -- EL
....
-----

JULY 8

30

HL

MAL

20

20

gc

" "'

10

1'1

13

15

11

13

15

19

17

E
40

I:~ 40'
J U L Y 13

it X

30

20,

iJ'xlJ

20

~ ~ -

10-

10

J U L Y 18

i",

30'

11

13

t (h)

15

17

19

11

13

15

17

19

t (h)

Fig. 5. (a) Canopy conductance (gs) scaled up by different methods from stomatal conductance
measurements in a maize field in Ottawa (1988) ( B A = b u l k average over the whole canopy,
T S = t o p sunlit leaves only, W = s u n l i t and shaded leaves weighted over the whole canopy,
EL = effective L, H L = sunlit and shaded leaves weighted over individual horizontal canopy layers, MAL = sunlit and shaded leaves weighted over multiple leaf angle classes in individual canopy layers) compared with surface conductance to water vapor transfer (go). (b) Canopy conductance (gs) scaled up by different methods (abbreviations are the same as in (a) from stomatal
conductance measurements in a maize field at Ottawa ( 1988 ) compared with surface conductance to water vapor transfer (go).

ESTIMATIONOF MAIZE CANOPY CONDUCTANCE

255

(b)

50

JULY 29
40.
i - ~

BA

....

TS

30.
**

EL

20.
......

HL
MAL

10.

gc
0

11

13

15

17

19

E
E

so I

o~ 50

'
40 t

, AUGUST 8
40

,o]

t ~

I t

30

_1

20

AUGUST 18

20

I
10

11

13
t (h)

15

17

19

7 ' ~ '1'1 ' 1'3 ' l g

' 1'7'1~

t (h)

in the season is the result of its capacity to better describe the radiation load
received by sunlit leaves. It is for this reason that it performs better on 8 and
13 July when an incomplete canopy leads to a larger fraction of L in the sun,
and late in the season (18 August) when lower sun elevations amplify the
difference between horizontal plane sampling and spherical leaf distribution
conditions. These results, therefore, confirm that sampling leafgs on a horizontal plane contributes to an overestimation Ofgc.

Leaf ternperature and water potential corrections

Corrections of gs for non-optimal leaf water potential were made only on
the three days when that parameter was measured (Fig. 6). Figure 7 shows
the effect of leaf temperature and leaf water potential corrections on gs when
eqn. (6) estimates for 13, 18 and 29 July were scaled up using the MAL

256

P. ROCHETTE ET AL.

BOTTOM PAF T OF CANOPY

0
.,4

"l I

-8

1
]

-16 .I

-12

- 2 0 - " ,

,.-.

".o
""
X

July 13

-16

- - - - July 18
SHADE - - - July 29
9

SUN
.

11 13 15 17 7 9 11 13 15 17 19
MIDDLE PART OF CANOPY

0
~.

ft.

~-4

,I ~ ' P ' "

-4

-J

< -8

-8

V"Y

5tu -12

"

-12

n- -16
uJ
SHADE

~ -2G

o
-4

-16
SUN
!

LL
_J

-20

11 13 15 17 7 9 11 13 15 17 1!
TOP PART OF CANOPY

I
\

-20

.-.

-4
-8

-8

i
-12

\-,~'~

-16

-12
-16

SHADE
"2C7 "9" "1 i "13 "15 "17" 7 "9" "1i "1:3 "15 "1~' "1!
t(h)

-2O

Fig. 6. Leaf water potential diurnal variation during 3 days of the 1988 growing season at three
levels in a maize canopy at Ottawa.

method. The leaf temperature was very close to the o p t i m u m on each day so
the correction for T~ m a d e insignificant modifications to eqn. (5) estimates.
The ~ corrections did not result in a better correspondence with go.
The curves on 18 July are o f particular interest. On that day, clear sky was
present until 12:00 EST, when a uniform cloud cover settled in. It can be seen
that g, obtained from measured g, values ( H L ) were close to go, especially
during the rapidly changing light conditions shortly after midday. The g, obtained from estimated g~ (eqn. (6) ) failed to follow the same curve. It there-

257

ESTIMATIONOF MAIZECANOPYCONDUCTANCE
3

20'
JULY 13

15

10

~.././

"-"\

'x

JULY 18

~"~

"~-~.~
12t /

"V '(i

\\

A
9

E
E

11

13 15 17 19

OI . . . . . . . . . . . .
7

9 11 13 15 17 19

t (h)

t (h)

w 40'
O~
35'

LEGEND

30'
25'

....
......

20'
15.

TI corrected
TI and LWP corrected
M e a s u r e d (HL)
gc

10'
5'

JU LY 29

11

13 15 17 19

t (h)
Fig. 7. Effect of correction for non-optimum leaf temperature (T~) and leaf water potential
(LWP) on canopy conductance estimates (gs) obtained by the multiple leaf angle classes layer
(MAL) scaling-up method (HL = canopy conductance obtained by scaling up stomatal conductance measurements made on a horizontal plane; go=surface conductance to water vapor
transfer)

fore seems that g~ responded to other stimuli than those used in eqn. (6), or
to the same stimuli but in a different way. These observations show that the
relationships between g~ and environmental parameters determined at high
Qp may not be valid in lower radiative conditions.

Shelter factor
Values ofgs obtained from gs measurements or estimates are useful for the
calculation of evaporation using the P e n m a n - M o n t e i t h equation. Models for
the estimation of latent heat transfer (Sinclair et al., 1976; Sellers et al., 1986 )
and dry deposition of trace gases (Baldocchi et al., 1987 ) include a canopy
conductance subroutine based on the scaling up of estimated leaf gs values.
Such a procedure could also be used with porometry measurements on plant
species for which insufficient information is available on the response ofgs to

258

P. ROCHETTEET AL.

TABLE 2
Daily values of the ratio of canopy conductance obtained by scaling up stomatal conductance (g~) to
that given by the Penman-Monteith equation (go) for different scaling-up methods (TS= top sunlit
leaves; HL = horizontal canopy layers; MAL = multiple leaf angle classes canopy layers )
Daya

Shelter factor

(gs/gc)

TS

HL

MAL

8 July
13 July
18 July
8 August
18 August

1.96
2.67
1.62
1.62
2.65

1.41
2.01
1.09
1.09
1.76

1.16
1.67
1.13
1.04
1.28

Mean

2.1

1.47

1.26

aJuly 29 was omitted because of wet soil conditions.

A 3.0

0 1.8

~ 1.2

~*

IIiu
a

alp

8
ull

W
~ 0.6
~ O.0

no

nmcl

II

10

100

ga,M / g s,T
Fig. 8. Relationship between the shelter factor (gs/gc) for the multiple leaf angle classes layer
scaling-up method (MAL) and the ratio of canopy aerodynamic conductance (ga,M) and the
stomatal conductance at the top of the canopy (gs,r).

the pertinent environmental variables. However, the results in this study have
shown that direct use ofgs obtained by the six methods tested is often unsatisfactory. Therefore, it is useful to find an empirical relationship between gs
and go. Shuttleworth ( 1976 ) suggested the use of a shelter factor (F) defined
here as

F~~g~'~
gc

(9)

where the subscript i refers to the scaling-up method. Mean shelter factor values over the growing season ranged from 1.3 for an assumption of spherical

ESTIMATION OF MAIZE CANOPY CONDUCTANCE

259

leaf distribution to 2.1 when only the sunlit leaves in the top 25% of the canopy were sampled (Table 2 ).
Finnigan and Raupach ( 1987 ) suggested that the g~/g~ ratio or shelter factor could be expressed as a function of the ratio of the aerodynamic and stomatal conductances of the leaves at the top of the canopy. Figure 8 shows the
relationship between the hourly values of the shelter factor (for the MAL
m e t h o d ) and the ratio of the canopy aerodynamic conductance (ga,M) over
g~ of the sunlit top leaves (gs,T). Data obtained when either the leaves or soil
surface were wet or when the sky was overcast were omitted. Scatter is large,
as expected when estimating g~ from 2E measurements, because small errors
in 2E can produce relatively larger errors in go. Nevertheless, we can identify
a tendency for the shelter factor to increase at higher ga,r~/g~,Tvalues in a way
similar to that predicted by the theoretical analysis of Finnigan and Raupach
(1987).
CONCLUSIONS

The diurnal and seasonal variation of maize gs was described for sunlit and
shaded leaves at three levels in the canopy. The diurnal variation in gs closely
followed photosynthetic photon flux density. The response Ofgs to photosynthetic photon flux density was dependent on the leaf level in the canopy and
growth stage.
Six different methods of scaling up from leaf stomatal conductance to canopy conductance were tested. Simpler methods limiting the sampling to the
top sunlit leaves on a horizontal position or involving the use of efficient L
(Szeicz and Long, 1969 ) gave the poorest results. The approach considering
a spherical distribution of leaf elements, where values of & are obtained from
the relationship between g~ and photosynthetic photon flux density, gave better estimates of gc. However, gc was still, on average, overestimated by 25%
and the use of a shelter factor is, therefore, recommended. The relatively poor
estimates of go obtained by the scaling up of leaf& in this study, tend to confirm that gc is not a purely physiological parameter, but also contains information about the net radiation load and aerodynamic conductance of individual leaves (Finnigan and Raupach, 1987 ).
ACKNOWLEDGEMENTS

The authors thank D. Balchin, W. Harrison, J. Desjardins, L. Houwing, B.

Dow, N. Crete, B. Boyd, D. Anderson and R. Verdon for their assistance in
field measurements and data analysis. This paper was written as a direct result of a workshop on stomatal resistance held at Pennsylvania State University, 10-13 April 1989. The workshop was made possible principally through
a grant from the National Science Foundation (BSR-8822164).

260

P. ROCHETTE ET AL.

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