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TRANSPIRATION AND LEAF TEMPERATURE

By DAVID M. GATESl
Missouri Botanical Garden, St. Loui, Misouri
INTRODUCTION
No literature of plant science ofers more confusion than the publica
tions concerned with transpiration and plant temperature. Transpiration
rate and leaf temperature are the result of the interaction of several simul
taneous environmental factors interacting with a leaf to a degree deter
mined by several plant properties. Hence the dependent variables, transpira
tion rate and leaf temperature, depend upon many independent variables of
climate and plant acting in various combinations at any single moment. The
investigator either has not recognized this inherent difculty, or complexity,
or if he did recognize it, did not know what to do about it. Therefore many
experiments which were thought to be defnitive were indefnite because one
or more variables were ignored in the context of the variables observed.
Very often good quantitative measurements were made of transpiration
rate, leaf temperature, air temperature, wind, and relative humidity and
then the relationships between dependent and independent variables were
not made quantitative for lack of a good analytical framework. In other
words, to put it simply and bluntly, the science of transpiration and leaf
temperature was not understood. It is hoped that this presentation will be a
step toward increased understanding of the interaction between transpira
tion, leaf temperatures, and environment.
ENERGY EXCHANGE
Environment is coupled to a leaf through the fow of energy. All plant
physiological processes, in fact all processes of life, require energy. The in
teraction of environment with a plant is through the fow of energy. There
is no other way. Energy delivered to the plant is converted to heat within
the plant, afecting the plant temperature, or energy is consumed through
photochemical and thermochemical events of the metabolic and physiologi
cal processes. All energy absorbed by a leaf must be accounted for and
hence the energy budget for a plant leaf must balance. The exchange of
energy by a plant leaf with its environment is written as follows:
Qa. = R C LE M
Each term is expressed in cal cm-2 min-l.
1.
Senior Fellow, Center for the Biolog of Natural Systes, Washington Univer
sity, which supported this reserch under PHS grant NO. 1 P. 10 ES 9139-01. Com
puter facilities were made available under NSF grant No. G 22296.
211
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Further
ANNUAL
REVIEWS
212 GATES
The radiant energy absorbed by the leaf is
Qaba'
A part of this, usually a
substantial part, is radiated by the leaf according to its surface temperature.
The radiation emitted is R. The air will cool the leaf by conducting heat
away if the air is cooler than the surface temperature of the leaf. The air
will deliver heat to the leaf by conduction if the air is warmer than the
leaf, and hence the negative sign in the second term on the right hand side
of Equation 1. There is always air movement around the leaf and the pro
cess of energy exchange between the leaf and the air is referred to as con
vection, indicated by C in Equation 1. In relatively still air there is free
convection or air movement simply caused by the temperature diferential
between leaf and air. In wind there is forced convection. This process has
been discussed by Raschke (47-49), by Gates (19-24), Wolpert (66), Wag
goner & Zelitch (65), Tibbals et al. (60), and by Knoerr & Gay (32) in
great detail. In addition, Gates & Benedict (25) have photographed the fow
of air around plants in order to demonstrate the character of convection.
Energy absorbed by a leaf goes into the evaporation of moisture which is
removed from the leaf by the combination of difusion and air movement.
L calories of energy per gram of liquid water are required to vaporize
water. Therefore, the energy consumed by transpiration is LE, where E is
the rate of transpiration in grams cm-2 min-1 If condensation of moisture
occurs on the leaf, heat is released to the leaf and the negative sign for the
third term on the right-hand side of Equation 1 applies. A small fraction of
the total energy
absorbed by a leaf goes into the metabolic processes and
this is indicated by M. If net photosynthesis occurs, incoming energy is con
sumed. If, however, stored energy is converted through respiration, a very
small amount of heating may occur. The most extreme case of respirational
heating may occur in the spathes of certain arums, see Beevers (6). Nor
mally, however, the energy consumed or contributed by the metabolic pro
cess within leaves is small and is negligible in the energy budget calculation.
A leaf receives radiation emitted by all surrounding surfaces, from
other leaves, the soil and rock surfaces, from clouds, and from the atmos
phere as well as from direct sunlight and scattered skylight. A leaf absorbs
a certain fraction of each incident stream of radiation, refects a fraction,
and transmits a fraction. The radiation absorbed by the two surfaces of the
leaf is -
2.
where _ _ _ and _ are the absorptances of the leaf surface to the
streams of direct sunlight, S; to the sLttered skylight, s; to the sunlight
and skylight refected to the leaf from the ground and plant surfaces,
reS + s); and to the infrared, thermal radiation emitted by the ground
surface, R!, and
by
the atmosphere, Ra' Each of the absorptances is wave
length-or frequency-dependent, as are also the intensities of the various
streams of radiation. The mean absorptance must be determined for the
specifc spectral composition of the incident radiation in a manner reported
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TRANSPIRATION AND LEF TEMPERATURE
213
by Gates et al. (28). Radiation is ubiquitous. At all times alI aerial parts of
plants are exposed to streams of thermal radiation from surrounding sur
faces, day or night, summer or winter. Sunlight and skylight are highly
variable and only present during the daytime, but thermal radiation is ever
present. The surface area of the leaf exposed to the sun as a point source
is Ai; to the skylight as a hemispherical, difuse source is A2; to the re
fected sunlight and skylight as a broad difuse source is Aa; and to the
hemispherical, difuse thermal radiation from the ground and atmosphere is
A4 The total surface area of the leaf is A. For a fat leaf A is twice the
surface area of one side of the leaf. Sometimes it is possible that Ai A2
Aa ~ A4 ~ A12. More frequently, most of the A's are diferent since a leaf
presents a different aspect to each source of radiation. All factors, such as
absorptances, radiation streams, and exposed surface areas, may change
with time.
The rate of water loss through transpiration by a leaf is determined by a
driving force and a resistance of the pathway. The driving force is es
sentially the diference, p, in concentration or pressure of water vapor
within the substomatal cavity of the mesophyll and of the free air beyond
the adhering boundary layer next to the leaf surface. The water vapor must
difuse from the sub-stomatal cavity through the stomatal channel and then
through the boundary layer into the free air where the moisture is carried
away by air movement. Hence the total resistance, T_ of the difusion path
way is given by an internal resistance, rz, made up of the sum of substomatal
plus stomatal resistances and an external resistance, T in the adhering air
layer. If there is a signifcant cuticular stream of moisture loss, this resistance
must be treated as a parallel pathway to the stomatal resistance. The rate of
transpiration per unit leaf area is given by
3.
where .p (T z) in g cm-2 min-i is the concentration of water vapor con
sidered at saturation at the leaf temperature, Tz, and
P
aCTa
)
in gm cm-2
min-i is the concentration of water vapor in the free air at the air tem
perature, Ta.
The thickness of the adhering boundary layer, and therefore the value of
the external difusion resistance, is a function of the leaf dimensions and
of the wind fow. Detailed laboratory determinations in a wind tunnel by our
group have permitted a new evaluation of this functional relationship. The
rate of transpiration is now written in the following form:
4.
where r.h. is the relative humidity of the air. .pa (Ta) is the saturated water
vapor concentration of the air at temperature, Ta, and k2 is a coefcient to
be discussed in a moment. The amount of transpiration is determined by the
energy available. The manner in which this manifests itself is through the
partitioning of energy in the energy budget and the adjustment of leaf tem-
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214
GATES
perature to a value which balances the energy budget relationship. The
leaf temperature determines the water vapor concentration or pressure with

in the leaf and therefore determines the driving force for transpiration.
Thus it is that the tran<piration rate and the leaf temperature are tied in
exorably together through the energy budget for the leaf.
The environmental factors which enter into the energy budget are, in
addition to the streams of radiation already described, the air temperature,
Ta in C, the relative humidity, r.h., and the wind speed V in cm sec-1. The
plant properties which enter into the energy budget are, in addition to the
absorptances and areas exposed, the emittance,

of the leaf surface to


thermal radiation, the leaf dimensions, D in cm in the direction of the wind
and W in cm transverse to the wind, and the internal difusion resistance
of the leaf, T_ in sec cm-1 The energy budget for H fat leaf is written:
where Tz in the first term on the right is in OK and
W D or W = D > 5 cm
WD OI W ..D5cm
kJ = 0.0100, k2 = 0.035
kt = 0.0162, k 0.026
The emittance,
is about 0.95 for most plants according to Gates & Tan
traporn (26). The coefcients kl and k2 and the powers assigned to D, W,
and V were determined by means of numerous wind tunnel experiments with
leaf models. Saturated blotting paper models were used to give the limiting
values corresponding to an internal difusion resistance of zero, and dry
blotting paper was used to simulate a leaf with infinite internal resistance.
Earlier we had always written two separate energy budget equations for free
convection in still air and one for forced convection in wind. Free convec
tion is considered to prevail at a wind speed of about 10 em see-1 Very
seldom in nature is the air movement less than 10 cm sec1 Further com
plications could be introduced into Equation 5 by introducing W in addition
to D in the convection term. The present equation is considered reasonably
accurate, but further improvements can be anticipated.
Because of the complexity of the dependence of leaf temperature and
transpiration rate on so many variables, it is impossible to visualize the
shape of the temperature or transpiration surface in this multidimensional
space. Hence many variables will be fxed and leaf temperature or transpir
ation rate examined as a function of one or two variables. The fgures pre
sented here are slices through this multidimensional space taken along the
dimension of a selected variable. Hence we shall have leaf temperature as a
function of the wind speed, or leaf temperature versus internal difusion re
sistance, or transpiration rate as a function of air temperature, or transpir
ation rate as a function of the wind speed, etc.
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TRANSPIRATION AND LEAF TEMPERATURE 215
V ARIATlON WITH WIND SPEED AND DIFFUSION RESISTANCE
Consider the relationship between transpiration rate and leaf temperature
for various amonts of wind movement from 10 cm sec-1 to 500 cm sec-1
and for various values of internal difusion resistance ranging from 0 to O
all other conditions fxed. Figure 1 gives transpiration rate versus leaf tem
perature for various wind speeds and for a leaf of dimensions 5 cm X 5 cm
which absorbs 1.2 cal cm-2 min-1 of radiation at an air temperature of 400 C
and a relative humidity of 50 per cent. These conditions are common in
warm, sunny, humid regions of the world during summer months at mid
day. Several interesting relationships are established by this graph. Follow
along a constant wind speed line and the infuence of changing difusion re
sistance as it af\t transpiration rate and leaf temperature is determined.
In still air (V ~ 10 cm sec-I) a change of internal resistancc from 20 sec
cm-1 to 2 sec cm-1 results in the transpiration rate changing froQ 17 X 10-5
to 64 X 10-5 gm cm-2 min-1 and the leaf temperature fro| 51 to 43 C. At
a wind speed of 100 cm sec-1 (2.2 mph) the changes are 11 X 10-5 to 70 X
10-5 and 45 to 4.70 C respectively. These are significant and substantial
diferences.
N ext it is of interest to note in Figure 1 the variations of transpiration
rate aMd leaf temperature when the wind speed varies and the internal dif
fusion resistance remains constant. At an internal resistance of 5 sec cm-1
or greater an increase of wind speed diminishes the leaf temperature and
the transpiration rate. At an internal difusion resistance of 2 sec cm-1 the
increase of wind speed increases transpiration rate and diminishes leaf tem
perature. At reSistance of 3 or 4 sec cm-1 there would be no change of
transpiration rate with increased wind speed. When the internal difusion
resistance is 0, case of wet blotting paper, the relationship between evapora
tion rate and surface temperature becomes very steep, and a change of wind
speed produces enormous changes of the evaporation rate while the temper
ature asymptotically approaches a lower limit, nearing the equivalent "wet
bulb" temperature of psychrometry. The region of the graph above and to
the right of the zero internal resistance line is one of "negative" internal
resistance which is equivalent to gutation or the forcing out of liquid under
pressure. The precise physical reality of any specifc position in this region
of the graph is unclear. At high wind speeds of 500 to 800 L sec-l or
greater the lines bend to the left and approach some nearly limiting max
imum evaporation rate well of the scale of the graph.
Figure 2 shows the same relationships as shown in Figure 1 except the
amount of radiation absorbed is now 0.8 cal cm-2 min-1 instead of 1.2 cal
cm-2 min-l. This graph might correspond to a warm, humid, cloudy summer
climate at midday. The change in the situation is now very striking. At in
ternal difusion resistances normally encountered with open stomates (r ~ 2
to 10 sec cm-1) an increase in wind speed always produces an increase in
transpiration rate and an increase in leaf temperature. At resistances be-
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216
30
220
E
200
E
W
M
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9
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"
GATES
Q
ab
s
" 1.
2 col cn2 mini. To" 40oC.
r.h." 50%. D=5cm. W"5cm.
Wet Bulb Pychrometric Temperature 30.
FIG. 1. Computed transpiration rate versus leaf temperature as a function of wind
speed and internal difusive resistance for the conditions indicated.
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80
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40
20
0
TRANSPIRATION AND LEAF TEMPERATURE
217
Qa
bs
=O.Scal cn
2 mini, T
a
=40C,
r.h.=50%, D=5cm, W=5cm,
Wet Bulb Psychrometric Temperature 30.6C
:
V;500
G
\
\
i
\
r= 2 sec cnl
LEAF
FIG. 2. Computed transpiration rate versus leaf temperature as a function of wind
speed and internal difusive resistance for the conditions indicated.
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218
GATES
tween 0 and 2 sec e-1 the increase of transpiration rate is enormous with
relatively small changes of leaf temperature with increasing wind speed.
The loss of water is always greater when the absorbed radiation is 1.2 cal
cm-2 min-1 (Fig. 1) than 0.8 cal cm-2 min-1 (Fig. 2) for the same value of
internal resistance.
At an air temperature of 200 C, elative humidity of 50 per cent, and
radiation absorbed of 1.2 cal cm-2 min-1 the efects of wid speed and dif
fusion resistance on transpiration rate and leaf temperature are shown in
Figure 3. These conditions of temperature and humidity would prevail for
comfortable clear spring days at mid latitudes or for summer days in high
mountainous regions. Leaves in bright sunshine, with resistances 2 sec cm-1
or greater, will have a slight decrease in transpiration rate and strong re
duction of leaf temperature with increasing wind speed. Under cloudy skies
with low amounts of absorbed radiation (about 0.6 cal cm-2 min-1) at an
air temperature of 200 C and r.h. = 50 per cent, an increase of wind speed
will increase transpiration rates and increase leaf temperature and the dia
gram would appear similar to Figure 2. The efects will be more pronounced
in drier air and less pronounced in more humid air.
.
It should be noted that the greatest infuence on transpiration and tem
perature is by the low wind speeds, under about 50 em see-1 or 1.1 mph.
These very low wind speeds are nearly always present in nature, and the
most frequently occurring changes of wind speed are from 4.7 cm sec-l (1
mph) to about 134.1 cm sec-1 (3 mph) or a little greater. Only very rarely is
nearly still air more stationary than 40 or SO cm sec-1 and for all practical
purposes free convection in still air corresponds to air movement of about
10 cm sec-1.
The infuence of wind movement at constant amount of absorbed radia
tion, but at diferent air temperatures and relative humidities, is shown in
Figure 4. Transpiration into dry air will increase with an increase of wind
speed. Transpiration into humid air will decrease with an increase of wind
speed if the amount of radiation absorbed by the leaf is large (1.2 cal cm-2
min-1). The same phenomenon is reported by Linacre (4) and earlier by
Raschke (48) in which an energy budget analysis was used which was simi
lar to the one presented here. When the amount of absorbed radiation i s
low (0.8 cal cm-2 min-1) then for moist ai r conditions a change of wind
speed will result in little change of water loss. For any amount of absorbed
radiation there are usually conditions during which there is no change in
transpiration rate with changes of wind speed. Ting & Loomis (62) re
ported the efect of wind and water loss from perforated septa, an experi
ment not completely analogous to the loss of water from leaves.
These efects are understood in the following manner. Recall that the
amount of water lost from the leaf by transpiration is determined by the
energy available and by a driving force created by a diference of vapor
pressure or water vapor concentration between the substomatal cavity and
free air beyond the adhering boundary layer. In addition, the rate of water
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180
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a
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"
TRANSPIRATION AND LEF TEMPERATURE
219
2 sec eml
Q
a
bs =1.2 c
a
l cm
2
mini.
r.h.= 50%. O=5cm,
Ta=20C,
W=5cm,
Wet Bu
l
b Psychrometric Temperature 14C
o
r=O
15
FIG. 3. Computed transpiration rate versus lef temperature as a function of wind
speed and internal difusive resistance for the conditions indicated.
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220 GATES
W, r= c sec eml
OO8 '~
LCr%
:
l"|
L" ZL%
ZL"L,
^ ?
-r =

WN bbOc B0Cl
Fm. 4. Computed trn . pran rate a a fuction of te wild speed fo Vl,OlS "if
tempertures ard retiv . h>1liditier for the fxed "ondtions given_
loss by the leaf is lc"trict<d by the resistance of the difulion pathway from
the sltbstomatal tgcu o\ hc leaf thru"gh tb< stomatal channel and then
th!Qugh the adhering boundary layer to the free ar beyond. The buundary
layer resistance is referrw to as the external resistance, and that arising
within the leaf as the internal resistance. The t<t.t difusion reslstallce of
the difusion pathway h the sum of the cxt.rnal and internal Iesistances.
An incease of wind sp<ed does two things wheI, considered at constant air
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.
TRANSPIRATION AND LEAF TEMPERATURE
221
temperature and relative humidity. It reduces the thickness of the adhering
boundary layer of air and thereby reduces the external resistance, and se
condly it changes the temperature of the leaf. The leaf temperature will in
crease with an increase of wind speed if the air temperature is warmer than
the leaf, which it usually is if the amount of absorbed radiation is low. The
water vapor pressure or concentration at saturation within the leaf will in
crease with increasing leaf temperature, and therefore the driving force for
transpiration will increase. The reduction of external resistance with in
creased wind speed and the simultaneous warming of the leaf caused by in
creased convectional transfer of heat to the leaf from the warm air will
each work to increase the rate of transpiration. If, however, the leaf is
warmer than the air at low wind speeds, which it often is when the amount
of absorbed radiation is high, as in full sunshine, then an increase of wind
speed wiIl cool the leaf by convection. The diminished leaf temperature will
diminish the water vapor pressure or concentration within the substomata1
cavity and thereby diminish the strength of the driving force, e.g., the
diference of vapor pressure between the substomatal cavity and the free
air. If the cooling of the leaf due to wind produces a greater relative
change of the driving force than the relative reduction in toial resistance
caused by the reduced external resistance, then the transpiration rate will
reduce with an increase of wind speed. Clearly, there are times when wind
causes a change of the boundary layer resistance which is just balanced b}
a change of the vapor pressure within the leaf, and the result is essentially
no change of the rate of water loss. Linacre (4) presented many of the
same arguments used here to explain the efect of wind and transpiration.
The infuence of leaf temperature and vapor pressure was discussed by Ru
felt, Jarvis & Jarvis (50).
The efects discussed above apply to a single leaf. What happens when
there is a change of wind speed throughout a plant canopy? Usually there
is an increased exchange of air within and without the canopy. During day
light morning hours it is likely that an increase of wind results in the
warmer air from outside the canopy mixing and raising the temperature of
the air within the canopy. Also it is likely that the air within the canopy
was more humid than the air outside the canopy, hence the mixing caused
by wind will produce somewhat drier air within the plant canopy. Many of
the canopy leaves are shaded and hence are in a radiation regime approxi
mating 0.8 cal cm-2 min-l. From Figure 4 one can see that wind bringing
warmer drier air into the canopy wiIl result in a substantial increase of
transpiration rate of the canopy. Only rarely will an increase of wind
within a plant canopy produce a measureable decrease of the loss of mois
ture within a canopy. If many leaves of a plant canopy are strongly sunlit,
as at midday, and are warmer than the air within the canopy, and the aiI
within the canopy is warmer than the air outside the canopy, then an in
crease of wind speed may result in a decrease of water usage by the can
opy.
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222
8
M
Q
z
o

zo
ec
9
30
a.
C
z

t
zo
|o
o
,
|o:a|:--|-|
V=IO em
,
-r'
r.h.=50%
D=5em
W=5cm
GATES
FIG. 5. Computed transpiration rate versus leaf temperature as a function of the air
temperature ad the interal difusive resistance for the conditions indicated.
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TRANSPIRATION AND LEF TEMPERATURE
223
A variation of wind speed in warm, humid, tropical-like environments is
likely not to produce a significant variation in the rate of water loss from
the vegetation. An increase of wind in warm, dry, desert-like environments
will result always in increased water loss from plants. An increase of wind
in cool, dry, or humid situations is likely to result in relatively little change
of water loss by plants.
VARIATION WITH AIR TEMPERATURE AND DIFFUSION RESISTANCE
(UNDER AND OVER AIR TEMPERATURE LEAVES)
Transpiration rate and leaf temperature as a function of the air tem
perature is shown in Figure S for nearly still air conditions, a relative
humidity of 50 per cent and absorbed radiation of 1.0 cal cm-2 min-l by a
leaf of 5 X 5 cm. An increase of air temperature increases transpiration rate
and leaf temperature for all values of internal difusion resistance. The lower
the internal resistance the more rapidly transpiration increases with air
temperature, and the leaf temperature increases somewhat less rapidly. It
clearly is seen when leaf temperature is above or below air temperature. In
nearly still air at 50 per cent relative humidity a leaf of dimension 5 cm X
5 em which absorbs 1.0 cal em-
2
min-1 of radiation will have temperatures
below air temperature only at high air temperatures (about 40 C or greater)
and at very low values of internal difusion resistance (less than about 2.0
sec cm-! ). With wind of 200 em see-l a similar diagram is shown in Figure 6.
Here a leaf of S cm X 5 L is cooler than the air if it has low difusion
resistance, but for somewhat lower air temperatures than for nearly still air
conditions.
The phenomenon of leaf temperature being under or over air temperature
when fully exposed to sunshine was reported by Lange (35) and Lange &
Lange (37) for leaves of many plants from Mauretania and the Costa Brava.
When the air temperature was 48 C the leaf temperatures were 38 C.
Lange also demonstrated the infuence of transpiration on leaf cooling by
excising these leaves and observing an immediate temperature increase from
38 C to 60 C in full sunlight. It is clear from Figure 5 that for Lange's
measurements the amount of radiation absorbed was high (approximately
1.2 cal cm-2 min-l), the wind speed was low and the internal difusion re
sistance of the attached leaf was very low (under 2 sec cm-l). Under these
circumstances calculation would agree with his observation. All variables
were not measured by Lange and therefore the observations cannot be
checked against calculation with precision. Lange (36) reported other mea
surements of transpiration and leaf temperatures under controlled conditions
for Phoenix dactylifera and Cucumis prophetarum.
Gates et al. (27) reported feld observations of Mimulus wherein leaf
tfmperatures were distinctly below air temperature in intense sunlight and
nearly still air conditions. When the air temperature was 18 to 20 C, the
temperatures of sunlit Mimulus leaves were 2S to 30 C. When the air tem
perature was 37 C, the sunlit leaves were from 30 to 35 C. Leaf tempera-
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22
4
140
120
40
2
0
Qabs = I. Oeo


V=2
0
em
s
ecl

r.h.
=50
%
0:5 em
W
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A
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/5
/0
35
F
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6.
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,
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,'"
1""
"
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'n
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,
'nn
"",
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and
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,
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"
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'ht
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'nr
t
,nn
rut
io
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TRANSPIRATION AND LEAF TEMPERATURE
225
ture equalled air temperature at 300 C. The relative humidity was very low
under these feld conditions, probably 20 per cent or lower, although it was
not measured. The
Q
ao8
was estimated from the following values for the day
of the measurements: S ~ 1.3 cal em' min'1 j ~ 0.15 cal cm'2 min'1 j
reS + s) ~ 0.25 cal em' min'!; Ra ~ 0.3 cal cm'2 min'! and Rg ~ 0.7 cal
cm'! 2 min'!; ai 02 = 03 0.55 and 04 0.95. The absorptivity for
Mimulus was reported by Gates et a1. (2). Hence, Qabs 0.94 cal cm'2 min'i.
The air movement was very little, probably between 10 and 50 em sec'1 and
at times up to 100 em sec'i. If the leaf dimension is taken as D ~ 2 em and
W ~ 5 em then Table I is computed from Equation 5 giving leaf tempera
ture and evaporation rate.
TABLE I
LEAF TEMPERATURE AND TRANSPIRATION RATE FOR Mimulus cardinalis AS A
FUNCTION OF WIND SPEED AND INTERNAL DIFFUSIVE RESISTANCE FOR A Qa.
OF ABOUT 0.94 LA L
7
1^
1
Wind speed cm sec-! 10 10 50 50 100 100
Leaf resistance sec cm-J 2 5 2 5 2 5
Leaf temperature, 0 C 30.1 32.7 29.4 31.0 29.3 31.0
Transpiration rate X 101
gm cm-2 min-1 54.0 30.8 62.7 30.0 64.1 30.2
It would appear from this that Mimulus cardinalis in the feld had an
internal difusion resistance of about 3.0 sec cm'l and a transpiration rate
in full sunlight of about 50 to 55 X 105 gm cm'2 min'!. A theoretical calcula
tion of the internal difusion resistance following the mCthod reported by
Lee & Gates (38) gave for full open stomates a value of 3.4 sec cm'i as
reported in Gates et a1. (27). At the time we made the observations we
could not calculate the energy budget with the precision permitted by the
current computer program. The agreement between the estimated internal
difusion resistance, based on the energy budget analysis for the tempera
ture at which leaf and air temperatures were equal, and the theoretically
calculated value is very good indeed.
Measurements made in a growth chamber by Gates et a!. (27) of leaf
temperature as a function of the air temperature gave the results shown in
Figure 7. It is seen that most of the plants became cooler than the air be
tween an air temperature of 30 to 40C and below these temperatures the
leaves were warmer than the air. From the energy budget the relationship
between leaf temperature and air temperature was calculated for conditions
approximating those of the growth chamber and the results are shown in
Figure 8. The calculations were at a constant Qabs ~ 0.8 cal cm,2 min'l
whereas in the growth chamber as the air temperature changed the amount
of radiation on the plant leaves changed slightly. Also as leaf temperature
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226

.
L
a:
::
t
a:
W
a.

L
..
l.
<
w
-
GATES
90
GROWTH CHAMBER
MIMULUS CARPINALIS (LOS TRANCOS)
80
o CRASSULA LACTEA
x OPUNTIA MONACANTHA
6 MIMULUS CARDINALIS (PRIEST'S GRADE)
o RHODODENDRON INDICUM (HORT. HYBRID)
70 MODERATE ILLUMINATION
8 MAY, 1964
GROWTH CHAMBER
60
+ AGAVE AMERICANA
' RAPHIOLEPIS OVATA
MIMULUS CARDINALIS (LOS TRANCOS)
* NERIUM OLEANDER
50
MODERATE ILLUMINATION
*
40

4.
.30
20
10
10 20
AIR TEMPERATURE, C
FIG. 7. Measured leaf temperature as a function of the air temperature for plants
i a growth chamber exposed to illumination of 1700 foot-candles at about 65 per cent
n:lative huidio.
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TRANSPIRATION AND LEAF TEMPERATURE
227
increased there was a tendency for the leaf internal resistance to drop, and
therefore at high air temperatures l ower leaf temperatures would result. Gates
et al. (27) reported an abrupt drop at leaf temperature of 410 C in total
leaf resistance from about 6 to 2 sec cm-1 The leaf resistances of 6 or
greater reported for Mimulus were at light intensities insufcient to open
stomates fully. It is clear by comparing Figure 7 and Figure 8 that Mimulus
had a low internal resistance and Opuntia, Crassula, A
g
ave, Nerium, and
50r
--------------
(
.40
I
a
:
<
a
I

30
L
t
I
<
L
.
Qabs=
0
.8
V=IO em sec
l
r.h.=65%
D=2em
W=5cm
30 40
AIR TEMPERATURE,C
50 60
FIG. 8. Computed leaf temperature versus air temperature for various internal difu
sive resistaces for the conditions indicated.
Rhododendron had substantially higher resistances. Above a resistance of
10 sec cm-1 the afect on leaf temperature is relatively small. A diagram
similar to these was presented by Linacre (39) when comparing leaf and
air temperatures for many plant species. He reports leaf temperatures tend
to equal air temperature at about 33 C. This cannot be universal but must
depend on radiation and other factors. It is interesting to note that Miller
& Saunders (42) reported temperatures of leaves of sorghum, corn, soy
beans, cowpeas, watermelon, pumpkin, and alfalfa to be very close to air
temperature in the vicinity of 30 C. They stated marked disagreement with
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228
GATES
the work of others reporting large diferences between air and leaf tempera
tures. It is entirely likely that Miller & Saunders were working close to the
crossover point between over and under-air temperature and that all in
vestigators obtained correct results.
VARIATION WITH LEAF DIMENSION AND DIFFUSION RESISTANCE
Leaf temperature and transpiration rate each are infuenced by the leaf
dimensions since the dimensions D along the direction of the wind fow, and
W across the direction of the wind afect both the convection and evapora
tion terms of the energy budget. The extent of this infuence is seen in Fig
ures 9, 10, and 11.
The behavior of transpiration rate per unit leaf area and leaf tempera
ture as a function of leaf dimension for semi-arid, sunny, summer day con
ditions is shown in Figures 9 and 10. The results are very striking. If a leaf
has a low internal difusion resistance, it will have a high rate of transpira
tion, total water consumption is enormous, and the amount of water used
per unit of photosynthesizing area is much greater for leaves of small di
mensions than it is for leaves of large dimensions. A change of leaf size for
low difusion resistance makes great diference in transpiration rate but
ver little diference in the leaf temperature, which is less than the air tem
perature.
If water is limiting, as it is in semi-arid regions, the plant leaves must
have relatively high difusion resistance to water vapor. It is clearly seen
in Figure 9 that for high resistance a change of leaf size produces a large
change of leaf temperature and a relatively small change of transpiration
rate. Small leaf size keeps leaf temperature near to air temperature. The
plants of arid or semi-arid regions tend to have small leaves or leaves which
are finely divided, often of a few millimeters. These plants usually are of
relatively dark coloration because of their thick, succulent character. There
fore the leaves do not transmit much radiation and their absorptance tends
to be particularly high, see Gates et aI. (28). Because of the intensity of
direct sunlight, refected sunlight, and intense radiant heat, the radiation
absorbed by a leaf may be 1.2 to 1.4 cal cm-z min-l. The question is often
raised: is there an adaptive advantage for a plant to possess small leaves
rather than large leaves from an environmental standpoint for hot, dry,
sunny conditions? The answer is clearly yes as seen from Figure 9. There
are two distinct advantages, namely reduced transpiration and reduced leaf
temperature. Although the air temperature is hot (4 to 50 C) at times in
these semi-arid regions, it is of considerable physiological advantage to the
plant to have leaf temperatures near to the air temperature rather than 5 to
15 C above air temperature. Many physiological processes, such as photo
synthesis, protoplasmic streaming, etc., begin to break down at temperatures
above 42 C or so [see Alexandrov (1)], and when leaf temperatures of
50 C or greater are reached, irreversible denaturation of plant proteins
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TRANSPIRATION AND LEAF TEMPERATURE
229
".'
'o
o ',1
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% "
.
N
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r",
30
Qa
b
s; 1.2 c
a
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2
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'
rh
.
=20%
l
V=IO em see
l
\
I
l


'
l
I
\
r=4
r
=
8
FIG. 9. Computed transpiration rate versus leaf temperature for various leaf dimen
sions and interal difusive resistances for the conditions indicated,
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230
40
T
E

3
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I
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a
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20
a
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z
c:
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= 20%
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a=40C
V=IOO em secl
r=IO em secl
GATES
Qabs
=
1.2
em2 mini
0 w
em em
I I 0
10 1
0
X
5
0 I
o
LEAF TEMPERATURE,oC
FIG. 10. Computed transpiration rate versus leaf temperature for various amounts of
absorbed radiation and various leaf dimensions for the conditions indicated.
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.
TRANSPIRATION AND LEA: F TEMPERATURE
231
occurs. The smallness of the leaves reduces the danger of leaf temperatures
rising unduly high, except perhaps for short periods when air temperatures
reach unusual maxima. Furthermore, if water becomes limiting, the stomates
will close, the internal difusion resistance will rise, the transpiration rate
will drop and the leaf temperature will rise. Once again small size (1 X 1 cm
or less) makes a signifcant diference in the leaf temperature compared
with the temperature of larger leaves when the internal difusion resistance
is large.
The efect of a change of radiation intensity and leaf size on transpiration
rate and leaf temperature is shown in Figure 10. Leaves of small size will
have relatively little variation of transpiration rate and leaf temperatures
with a change in the amount of absorbed radiation. Variations of the
amount of absorbed radiation will occur from sun to shade, for changes of
angle between leaf and the solar rays, for cloudy to sunny conditions, or
from night to day. Leaves of large size undergo enormous changes of trans
piration rate and changes of leaf temperatures with variations of the amount
of absorbed radiation. Once again the advantage of having leaf temperature
tightly coupled to air temperature dictates small leaf size. Some succulent
plants tend to keep stomates open at night as discussed in detail later in this
paper. Although we cannot explore the problem in complete detail here, it is
evident from Figure 10 that at Qabs ~ 0.6 cal cm-2 min-I increased size im
plies reduced transpiration rate per unit area and also reduced leaf tempera
ture. So perhaps at night when certain succulents assimilate CO
2
there are
adaptive advantages to be found here.
In hot, humid climates with ample water available for transpiration, a
low internal difusion resistance is possible and leaf temperature will tend
to be close to air temperature. This is shown in Figure 11. Low internal
difusion resistance must go with large leaf size to keep the leaves from
becoming too warm. It would seem that ample water implies ample growth
under warm, humid, sunny conditions providing the leaves have low inter
nal resistance. A low resistance also implies better uptake of CO2 for assi
milation, which is also necessary for ample growth.
TRANSPIRATION-TEMPERATURE INVERSION
Of particular interest is the transpiration-temperature inversion point
which occurs when leaf temperature equals the air temperature as seen in
Figures 9 and 11 and also in Figures 1, 2, and 3. Below this point, all leaves
are above air temperature and leaf temperature increases with an increase
of leaf size. Above the "inversion" point, leaf temperature is always below
air temperature and transpiration rate per unit leaf area decreases with an
increase of leaf size. Above the "inversion" point a change of leaf size pro
duces relatively little change of leaf temperature when the radiation ab
sorbed by the leaf is high, e.g., 1.0 or 1.2 cal cm-2 min-I. The "inversion"
point drops along the line of leaf temperature equal to air temperature and
is zero at a point where there is no longer sufcient energy available to pro-
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232
I
'f
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0
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a:
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..
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.. "
'
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Ta=40C
'
.
0 ,
\
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h.=80%
'
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%
,
,
I
V =100 em
,
-rI
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.
.
.
20
I
10
r
=
'.
r=50
.
.
.
. r=
O
0-
LEAF TEMPERATURE,C
FIG. 11. Computed transpiration rate versus leaf temperature for various leaf dimen
sions and interal difusive resistances for the conditions indicated.
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TRANSPIRATION AND LEAF TEMPERATURE 233
duce evaporation of water. In faLt, if the amount of radiation absorbed is
very low and the air is relatively moist, condensation may occur.
The inversion point becomes a surface when it i s thought of in the con
text of all variables, e.g., radiation, wind speed, air temperature, and rela
tive humidity. In the vicinity of the inversion point, a change of leaf size
produces almost no change of leaf temperature nor of transpiration rate. It
is possible, for ample water available, that plants tend to select internal dif
fusion resistances according to the range of environmental conditions oc
curring most frequently such that they get as close to the inversion point,
line, or surface, as possible and thereby reduce the variations of transpira
tion and temperature caused by variations of leaf size. It wilJ take a great
deal of careful observation and measurement to prove or disprove such a
hypothesis.
TRANSPIRATION AND ITS TEMPERATURE AFFECT
If the plant physi ologist ever has had a doubt concerning the value to
the plant of transpirational cooling as it affects leaf temperature, this gross
misconception should be dispelled now once and for all. As we have seen,
there are conditions under which variations of transpiration rate may make
relatively little diference in leaf temperature. However, often the ability
to transpire will make a substantial diference in leaf temperature when
the heat load on a leaf is large and thermal death is likely to ensue if leaf
temperatures rise too high. The efect of changing interal difusion re
sistance to water vapor by opening or closing of stomates is dramatically
seen in most of the graphs presented here. From Figure 8 it is seen that a
leaf of dimension 10 em X 10 em transpiring 7.1 X 10-4 g cm-2 minot is
10 C cooler than a nontranspiring leaf when the air temperature is 40 C
and the other conditions consistent with Figure 8 pertain. Similar compari
sons can be made for any other selected conditions.
These predictions are borne out by Slatyer & Bierhuizen (57) where the
use of transpiration suppressants increHsed leaf temperature by as much as
9 C and by the splendid work of Martin (41 ) where a transpiration rate
of 16.6 X 10-5 g cm- min-1 produced a leaf temperature depression of
about 8 C for H elianthus annus and Ambrosia trifda. The work by Mar
tin came the closest of any up to that time to giving a careful quantitative
analysis of the role of convection and transpiration in producing tempera
ture efects in l eaves. Even so, Martin was not able to give the exact analy
sis of the energy budget of a leaf, in large part due to the fact that he did
not know the amount of radiation absorbed by the leaves used in his experi
ments. His estimates of the absorptivity of leaves were apparently very
wrong because of radiation errors with the thermocouples he used as radi
ometers. Linacre (4) discusses the relationship between transpiration rate
and leaf temperature tlsing the xteral boundary layer resistance as the
indepCndent variable.
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234 GATES
Curtis ( 12-14) , Curtis & Clum ( 15 ) , and Clum ( 10) believed they had
shown the relative unimportance of transpirational cooling in plants. Shull
( 54, 55) , Eaton & Belden ( 16) , Arthur & Stewart ( 2) and Clements (9)
were certain that transpiration produced considerable cooling of leaves. The
utter confusion and lack of agreement among these earlier investigators
was largely a lack of good physical theory applied to the problem of tran
spiration and leaf temperature. As long as they omitted a sound analytical
foundation from their discussion they could not hope to sort out their
differences.
Many of the earlier conclusions were confused further when leaves
were coated with vaseline or enclosed in cellophane and transpiration mea
sured. When this was done, the radiation absorbed and emitted by the leaf
was changed drastically, and the energy budget of the leaf was diferent
from the natural leaf. Without the use of physical theory and careful en
ergy budget analysis, comparison between the natural leaf and the modified
leaf was invalid.
Other workers such as Waggoner & Shaw ( 64) and Konis (33) ob
tained good temperature measurements of leaves and were able to point out
the importance of energy budget considerations. Yet they did not work out
a quantitative energy budget evaluation nor did they give specifc transpira
tion rates as related to changes in leaf tempertaure. Tanner ( 59) demon
strates that observations of plant temperatures are a possible index of water
relations for various corps. Audus & Cheetham (3) showed that ethereal
oils on leaves did not infuence significantly the transpiration rates nor the
leaf temperatures. Bregetova & Popova ( 7) found that for some heat unsta
ble plants, e.g., paspalum, cotton, corn, and tomato, the leaf temperature to
air temperature gradient is lowered in the warmest part of the day com
pared with the behavior of heat-stable grape leaves for which the gradient
increases in midday.
TRANSPIRATION AS A DIFFUSION PROCESS
The transport of water vapor from within a leaf to the free air beyond
the leaf is a difusion process with the water molecules travelling from a
region of greater water vapor concentration to a region of lesser water
vapor concentration. The pathway for difusion of water vapor from a leaf
is primarily from the mesophyll cell walls lining the substomatal cavity
through the stomates and then through a boundary layer adjacent to the
leaf surface to the free air beyond the boundary layer. A parallel pathway
is directly through the leaf cuticle and through the boundary layer. When
the stomates are open, far more water vapor escapes from the leaf along
the stomatal pathway than through the cuticle. The first detailed discussion
of the difusion of gases through stomates was the classical paper by Brown
& Escombe ( 8) in 1900. The earlier work in physics concerning gaseous
difusion is referred to in this paper and in particular Stefan's ( 58) deriva
tion of the law for difusion. Stefan is often misquoted as predicting diame-
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TRANS'PIRATION AND LEAF TEMPERATURE
235
ter proportionality for evaporation through a tube from a wet surface. Ac
tually Stefan's formulation had nothing to do with difusion through a tube.
The formulation of difusion theory most traditionally used by plant
scientists is that utilizing circuit theory. This was recently criticized by
Cooke ( 11) who has suggested substituting a feld theory approach for the
circuit theory approach. Since nearly all work concerning transpiration uses
the circuit theory terminology, this approach is used here. Lee & Gates (38)
have reviewed the history of the subject of difusion theory and utilized a
numerical approach suggested by Bange (4) for computing the difusion re
sistance of stomates. Such abused topics as the so-called "diameter law" for
difusion through a pore are discussed critically by Lee & Gates ( 38).
TABLE II
RESISTANCES TO THE DIFFUSIVE TRANSFER OF WATER VAPOR THROUGH THE
STOMATES, r .. 2M CUTICLE, r" AND THE BOUNDARY LAYER, f"
Plant r. r, r ra r
Populus tremula 2.3 2.3 0.6 2.9
Betula verrucosa 1 . 2 70 1 .2 0.8 2.0
Quercus robur
7 . 1 265 6.9 0.8 7.7
16.2 315 15 . 4 0.9 16.3
Acer platanoies 5.0 82 5.3 0.8 6 . 2
Circaea lutetiana 12.4 90 10.9 0.5 1 1.4
Lamium galeobdolon 10 . 0 33.0 7 .7 0.8 8.5
Helianthus annuus 0 . 4 0.4 0.5 1.0
Gossypium hirsutum 1 . 0 32.3 1 .0 2.1 3.1
The total leaf resistance is given by r/ =1,1./(r.+,,) and the total resistance of
the difusion pathway is given by r=rZ +ra. Resistanc units are in sec cm-i Taken
from Holmgren, Jarvis & Jarvis (29) except for cotton which is from Slatyer &
Bierhuizen (57).
Many investigators have used these formulations of resistance for leaves
to interpret experimental data, see Slayter & Bierhuizen ( 57) , Sayre ( 5 1 ) ,
Sierp & Seybold (56), Huber (30) , and Ting & Loomis (61) . For most
plants the boundary layer resistance is smaller than the internal resistance of
the leaf [see Miller & Gates (43) ] , but for agricultural plants and some non
cultivated plants [see Monteith (4) and Holmgren, Jarvis & Jarvis ( 29) ] ,
the interal leaf resistance is very low and the boundary layer resistance is
the larger of the two. The boundary layer resistance depends on the depth of
the boundary layer of adhering air, which in turn depends upon the leaf size,
shape, and orientation, upon the temperature diferential between leaf and
air, and upon wind speed. The boundary layer resistance becomes negligible
at wind speeds greater than 2 mph ( 89.4 cm sec-I) . Table II gives measured
values of T_ T_ and T for water vapor. The leaf resistance, T_ is determined
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236
GATES
TABLE I I I
INTERNAL LEA DIFFUSIVE RSISTANCES, 7] IN SEC CM-1 FOR
VARIOUS PANT LEAVES
Plant
Turnip, sugar beet
Barley
Tomato and Bean
Wheat
Zebrina pendula
Kaanchoe blossjeliana
Populus tremuloides (wet site)
Populus tremuloies (dry site)
Populus gradientata
A mmophia breveligulata
Arctostaphylos uva-ursi
Betula papyriera
Chamaedaphnt: calyculata
Acer rub rum
Vii
Quercus rubra
Pinus strobus
Pinus resinosa
Picea mariana
Thuja occientalis
Pteriium aquiinum
1.5- 1. 7
1 . 0- 2. 0
2 . 3- 3 . 3
0 . 2- 2 . 4
1 .08
1.20
2.0- 10.0
2.5- 2.8
4.0- 5.0
4. 0- 10. 0
3.0- 5.0
5 . 0- 6. 0
6. 0- 8. 0
9 . 0- 17 . 0
1 1.0- 20.0
1 1.0- 18.0
20.0- 29.0
32.0- 50.0
20.0- 60.0
45.0
70.0-200.0
150.0-400.0
Investigator
Gaastra (18)
Armstrong (Unpublished)
Kuiper (34)
Penman and Schofeld (45)
Bange (4)
Bavel, von (5)
Ting, Thompson and Dugger (63)
Miller and Gates (43)
Miller and Gates (43)
Miller and Gates (43)
Miller and Gates (43)
Miller and Gates (43)
MilleI and Gates (43)
Miller and Gates (43)
Miller and Gates (43)
Miller and Gates (43)
Miller and Gates (43)
Miller and Gates (43)
Miller and GHtes (43)
Miller and Gates (43)
Miller and Gates (43)
Miller and Gates (43)
by the stomatal resistance, T_ in parallel with a cuticular resistance, Tg-
Hence,
l/n = l/r. + l/r.
6.
Generally for open stomates Tg > > ? and T_ m ? However, with
stomates closed T_ ~ and T
[
~ T_. For many species with the stomates open
T_ and T_ are comparable in magnitude and both must be considered when
computing 1,.
DIFFUSIVE RESISTANCES
It is important to understand the values of the difusion resistances to
the extent reported in the literature. Table II lists values of
Tg
* [ ' @
and T
determined by Holmgren, Jarvis & Jarvis (29) for many plant species and
for cotton by Slatyer & Bierhuizen (57) . In Table III are listed values of
leaf resistance for many plants measured by various investigators. The
agricultural crops generally have the lowest values (0.2 to 3.3 sec cm-1 ) of
l eaf resistances, although there are certainly exceptions such as H elianthus
annus with a value of 0.4 sec cm-I. Many broad, deciduous leaves have values
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TRANSPIRATION AND LEF TEMPERATURE
237
of difusive resistances from about 5.0 to 20.0, pine needles from 20.0 to 60.0,
Thuja 70.0 to 200.0 and bracken fern, Pteridium aquilinum 150.0 to 400.0 sec
cm-l, Many more measuremnts need to be taken of nonagricultural plants
in order to get a better understanding of their water usage.
Although most plants assimilate during the daylight hours and have
stomates open in light, certain plants are unique in that assimilation takes
place in the dark according to Ranson & Thomas ( 4) and have greater sto
matal opening at night than in the daytime as reported by Shreve (53) . Ekern
( 17) reported measurements indicating that pineapple may lose more water
at night than during the daytime as also reported by Joshi, Boyer & Kramer
(31 ) . Recently Ting, Thompson & Dugger ( 63) reviewed the literature
concerning water loss from succulent plants and reported a thermoperiod
efect with the stomatal opening of Kalanchoe blossfeldiana. The diurnal
change of leaf resistance followed a normal pattern, low in light and higher
in darkness, when the plants were pretreated with cool thermoperiods or with
thermoperiods having little diurnal temperature variation. Large diurnal
temperature changes resulted in apparent nocturnal stomatal opening. In
actual fact it seemed that day stomatal closing occurred rather than com
plete nocturnal opening. The minimum average resistance in light for short
day plants of Kaanchoe was 1 1.6 sec cm-1 and in the dark was 58 sec cm-1
and for long day plants 1 1.5 sec cm-1 in the light and 72 sec cm-1 in the dark.
With increasing daytime temperatures, the difusion resistance in light
increased, and with increasing daytime temperatures, the difusion resistance
at night appeared to decrease. The ratio of nighttime to daytime resistances
was about 2.5 to 1 .0 at day temperatures from 15 to 23 C, while above 23 C
the daytime resistance exceeded nighttime resistance by as much as 3.5 to
1 .0. The nighttime resistance did not drop much in absolute value with high
daytime temperatures but the daytime resistance became unusually high.
Scarth (52) found that high temperatures (38 to 4 C) helped to increase
stomatal opening in light but tended to prevent their closure in darkness, and
that low temperatures (0 to 8 C) prevented stomatal opening in light.
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