Agricultural and Forest Meteorology, 42 (1988) 121 133 121

Elsevier Science Publishers B.V., Amsterdam Printed in The Netherlands

LEAF TEMPERATURES DURING RADIATION FROST

P A R T I. O B S E R V A T I O N S

R. LEUNING and K.W. CREMER

CSIRO Division of Forest Research, Box 4008, Queen Victoria Terrace, Canberra, A.C.T., 2600
(Australia)
(Received March 23, 1987; revision accepted June 2, 1987)

ABSTRACT

Leuning, R. and Cremer, K.W., 1987. Leaf temperatures during radiation frost. Part I. Observa-
tions. Agric. For. Meteorol., 42: 121-133.

Temperatures attained by leaves in natural environments are significantly influenced by

microhabitat and morphology of the plant. Effects of leaf size, height above ground and the type
of underlying surface on leaf temperatures of Eucalyptus pauciflora and E. viminalis seedlings
during nights of radiation frost were investigated.
Temperatures of leaves exposed to clear skies after sunset were 1-3C lower than those of the
adjacent air at all heights except ground level. Intensity and duration of sub-freezing temperatures
of leaves diminished with increasing height above ground. Temperatures at the centre of larger E.
pauciflora leaves were 0.5-1.5C lower than for smaller E. viminalis leaves, but tip temperatures of
the former were similar to those at the centre of E. viminalis. Leaf temperatures of E. pauciflora
were ~ 0.5-1.5C lower when held horizontally compared to vertical leaves at the same height. The
corresponding change in E. viminalis was between 0.2 and 0.6C. At each height, leaf temperatures
above grass were lower than those above dry soil and these were lower than those above moist soil.
Frequency distributions relating screen temperatures to those of leaves of isolated seedlings
were determined.

INTRODUCTION

T h e r e is a n e x t e n s i v e l i t e r a t u r e o n t h e r e s p o n s e of p l a n t s to f r e e z i n g tem-
p e r a t u r e i n t h e l a b o r a t o r y (e.g., S t e p o n k u s , 1978; L e v i t t , 1980) w h i c h s h o w s
t h a t b i o l o g i c a l r e s p o n s e to l o w t e m p e r a t u r e s m a y d e p e n d o n t h e d u r a t i o n a n d
i n t e n s i t y of c o l d c o n d i t i o n s a n d o n c r i t i c a l t e m p e r a t u r e t h r e s h o l d s . P h y s i c a l
factors c o n t r o l l i n g t e m p e r a t u r e s a t t a i n e d by p l a n t o r g a n s d u r i n g r a d i a t i o n
f r o s t s i n t h e o p e n w a r r a n t c l o s e a t t e n t i o n w h e n a t t e m p t s a r e m a d e to r e l a t e
l a b o r a t o r y r e s u l t s t o t h e field. T h e p r e s e n t p a p e r is c o n c e r n e d w i t h l e a f e n e r g y
balances and how tissue arrives at particular temperatures.
P l a n t r e s p o n s e t o f r o s t is o f t e n c o n s i d e r e d i n r e l a t i o n to a i r t e m p e r a t u r e s
m e a s u r e d i n a m e t e o r o l o g i c a l s c r e e n o r c l o s e to t h e s u r f a c e o f s h o r t g r a s s u s i n g
a b u l b - t h e r m o m e t e r (e.g., C a n n e l l , 1984). H o w e v e r , l e a f a n d b u d t e m p e r a t u r e s
m a y differ s i g n i f i c a n t l y f r o m b o t h t h e s e t e m p e r a t u r e s , d e p e n d i n g o n t h e
prevailing atmospheric conditions and on variations in plant microhabitat,

0168-1923/88/$03.50 1988 Elsevier Science Publishers B.V.

122

such as height above ground and the extent of exposure to night sky and to
surrounding surfaces. For example, Nunez and Sander (1982) found that air
temperatures at night were 1 2C warmer in a shelterwood of Eucalyptus
delegatensis compared with an adjacent clear-felled area. The shelterwood
provided a more favourable environment for seedling survival because there
was less loss of thermal radiation from the plants to the sky than occurred in
the open area.
Ludlow and Fisher (1976) showed t hat the amount of leaf death of Macrop-
tilium atropurpureum was directly correlated with the quantity of leaf litter
surrounding the plants. They suggested t hat litter impedes the flux of heat into
and from the soil, resulting in lower surface temperatures, lower leaf tem-
peratures and greater frost damage to leaves over litter compared to those
above bare soil. The apparent lower sensitivity to frost by seedlings relative to
mature plants was correctly attributed by Ludlow and Fisher (1976) to smaller
quantities of litter beneath seedlings compared to mature plants r a t h e r than to
inherent biological differences in their frost susceptibility.
Failure to appreciate physical influences on plant temperatures may
confound the int er pr et at i on of the biological response of plant organs
subjected to frost in the field.
Field data are presented to illustrate the influence of leaf size, leaf orienta-
tion and height above ground and the effect of the type of underlying surface
on leaf temperatures of young seedlings during nights of radiation frost.
Since the only widely available data on frost temperatures are screen
minima, frequency distributions relating screen and leaf temperatures at
various heights are also presented.
A detailed the or y for predicting leaf-to-air temperature differences as a
function of leaf size and temperatures of the air, screen and underlying surface
is presented in a companion paper (Leuning, 1987).

METHODS
Experimental data were collected in the meteorological enclosure at
Yarralumla, Canberra. The site had a slight slope ( ~ 2.5 ) and was covered in
dense grass ~ 2 5 m m high. There were 18-m tall trees 50m south and low
buildings 80m west of the experimental area.
Leaf temperatures were measured on eight nights by inserting a copper-
constantan thermocouple junction into the abaxial epidermis at the centre of
healthy E. pauciflora and E. viminalis leaves. Temperatures near the tips of E.
pauciflora leaves were also measured. The thermocouple wires were 75 ttm in
diameter and they were taped along the abaxial surface of the leaves to
minimize thermal conduction from the surrounding air along the wires to the
sensing junction. Reference junctions of the thermocouples were attached to a
reference platinum resistance t her m om e t e r placed in a dewar flask filled with
water.
Leaves of both species were held horizontally at heights of 0, 40, 100 and
1000 mm and vertically with their midribs at 40 and 100 mm above grass. Leaves
 123

at 0 mm were in c o n t a c t with grass tips. I n d i v i d u a l leaves were displayed so

t h e y were well exposed to b o t h the sky and the ground. Air t e m p e r a t u r e s were
m e a s u r e d at the same h e i g h t s using unshielded, u n a s p i r a t e d 75-pm copper-
c o n s t a n t a n t h e r m o c o u p l e s . A t h e r m o c o u p l e j u n c t i o n was also placed in a
n e a r b y s t a n d a r d m e t e o r o l o g i c a l screen. C a l c u l a t i o n of r a d i a t i o n e r r o r s on
spheres 150 pm in d i a m e t e r i n d i c a t e d t h a t air t e m p e r a t u r e s were m e a s u r e d to
+_0.05C.
A second set of t e m p e r a t u r e m e a s u r e m e n t s was t a k e n on t h r e e nights using
leaves of b o t h species held h o r i z o n t a l l y at 5, 40 and 100 mm above grass and
o v e r 1.2 1.2-m plots of moist and dry b a r e soil. F o r these m e a s u r e m e n t s , the
j u n c t i o n s were a t t a c h e d to the u n d e r s i d e of the leaves using porous tape
(Leucopor). Rain was excluded from the dry soil plot and the moist a r e a was
w a t e r e d r e g u l a r l y . M o i s t u r e c o n t e n t of the soil n e a r the surface was
determined gravimetrically.
M e a n s and s t a n d a r d d e v i a t i o n s of the leaf width m e a s u r e d at half-length
were 48 + 8 m m (n = 17) for E. pauciflora and 10 + 2 m m (n = 17) for E.
viminalis. Seedlings ~ 1 y e a r old and 1.2 m tall were growing in pots t h a t were
b u r i e d to p r o t e c t t h e i r roots from freezing.
A s c a n n e r - v o l t m e t e r ( H e w l e t t P a c k a r d HP3421A) c o n t r o l l e d by a small
c o m p u t e r (HP 41CV) was used to m e a s u r e the electrical signals e v e r y 30 s and

9 "" ".. lO00

~_100""....
"...:

E
\ '..':.
\ - .:... ..
,... ..- .

18 22 %0,:'%,2 6

t (h}

-4

Fig. 1. T i m e s e r i e s of air t e m p e r a t u r e s m e a s u r e d a t h e i g h t s (z) of 0, 40, 100 a n d 1000 m m above a

field of s h o r t g r a s s on 20-21 J u l y 1984. U p a r r o w s i n d i c a t e periods of w a r m i n g o b s e r v e d at all levels,
d o w n a r r o w s i n d i c a t e w a r m i n g at s o m e levels n o t o b s e r v e d at others. {Sunset 17:05h, s u n r i s e
07:05 h.)
124
provide 15-min averages of the temperatures. Accuracy of the measurements is
estimated to be __0.1C.

RESULTS AND DISCUSSION

Air temperatures

Typical time series for air temperatures at heights of 0, 40, 100 and 1000 mm
are shown in Fig. 1. Air temperatures measured in the screen (at 1.5 m) were
within 0.1C of those measured at 1000mm. Increasing air temperature with
height indicates stable stratification of the surface boundary layer. Although
there was a net cooling of the air each night, intermittent periods of warming
were occasionally observed simultaneously at all levels (e.g., at up arrows in
Fig. 1). Relatively large air motions were responsible for these fluctuations
since their influence persisted for periods greater than the 15-min averaging
time. On other occasions, fluctuations at one height were not observed at other
levels (e.g., at down arrows in Fig. 1), reflecting the highly intermittent and
discontinuous nature of air movements during stable stratification (e.g.,
Ogawa et al., 1985). The onset of fog at ~ 02.30 h caused all air temperatures to
converge.
Andrd and Mahrt (1982) stated that the surface inversion layer develops by:
(1) an upward extension by turbulence of cooling of air in contact with the
surface; (2) clear-air cooling of the air itself through a net emission of radiation
by molecules of water vapour, carbon dioxide and ozone and by aerosols; and
(3) advection of cold air from the surroundings. The equation for conservation
of heat in an infinitesimally thin layer of air during an averaging period At may
be expressed as:
oT o - - ~ ~R
- (w'T') O. AT (1)
t ~z paCp ~Z
where T is the mean temperature, Pa and Cp are the density and specific heat of
air, w'T' is the mean kinematic heat flux density, R the net long-wave radiative
flux density. The temperature advection rate is given by the last term in eq. 1,
in which 5 is the mean velocity vector. Following standard nomenclature, w'
and ~ represent instantaneous departures from mean values fi) and ~, respec-
tively. Radiative flux divergence (OR/~z) near the ground results predominant-
ly from gradients in temperature and in water vapour concentration.
Average rates of temperature change for six nights at heights of 0, 40, 100
and 1000 mm were - 4.3, - 3.2, - 2.6 and - 1.8C h 1for the hour around sunset.
After the initial rapid drop in temperature, mean cooling rates decreased
steadily to ~ - 0 . 5 C h -1 until the temperature minima were reached. Funk
(1960) measured cooling rates and radiative flux divergences directly and found
t h a t ~T/~t was often smaller in magnitude than that caused by ~R/~z. Equation
1 implies that divergence of turbulent heat flux and advection contributed to
warming from above of the air near the surface. This provides a satisfactory
 125

~ ~ _ ~ ,~_~
100(3

800

600
N

400

200

lO0

--4 -2 0 " 2 "4 - 6 " " I0- 12

T (C)

Fig. 2. Profiles of air t e m p e r a t u r e s m e a s u r e d on 20~21 J u l y 1984 s h o w i n g the development of the

surface inversion. The end-time of 15-min a v e r a g e s is s h o w n above each profile.

explanation for the relatively low cooling rates observed for most of the night
(Fig. 1). However, initial cooling rates of the air near sunset were larger than
can be expected from radiative cooling alone (Funk, 1960; Coantic and Seguin,
1971) and turbulent exchange of heat at the surface was important for cooling
the air during this period.
A series of profiles of air temperatures showing the development of the
surface inversion layer is presented in Fig. 2. Air at the ground was warmer
than air at higher levels until ~ 1.5 h before sunset, but then decreased rapidly.
Temperature gradients reversed first at the ground and then later for the whole
profile. The magnitude of the gradients also increased markedly until ~ 1 h
after sunset and then remained similar until the onset of fog at ~ 02.30h.
Similar results were obtained on other calm nights. A local temperature
maximum at 40 mm was observed on most evenings during the transition from
lapse to stable profiles (Fig. 2), showing development of the temperature
inversion from the surface upwards. However, the minima found by Lake (1956)
and Oke (1970) at heights above, rather t h a n at the ground surface, were never
observed in this study.
Grass swards or litter layers are relatively poor conductors of heat. Hence
air in contact with these surfaces cools to a lower temperature t h a n air in
contact with a better thermal conductor such as moist soil. This is illustrated
in Fig. 3, which shows air temperatures measured at 5 mm above short grass
126

x\~xx~Z'., wet soil . ///

\\xx_ xx- '" / ~~ ...
ry "-... .,
< grass ,,,, soil "'"'.... .:

I
18 20 22 0 " - - . ~ "2,~ ,~/ 4 6 8
t (h)

-2 t
fog

Fig. 3. Time series of air temperatures measured at 5 mm above 25-ramtall grass, dry soil (0g = 0.10)
and moist soil (0g = 0.26) on 15-16 August 1984. Onset of fog is marked. (Sunset 17.30h, sunrise
06.45 h.)

a n d o v e r d r y a n d m o i s t soils w i t h g r a v i m e t r i c m o i s t u r e c o n t e n t s (Og)of 0.10 a n d

0.26 g g 1, r e s p e c t i v e l y . A i r t e m p e r a t u r e s a b o v e g r a s s w e r e ~ IC less t h a n o v e r
dry soil a n d ~ 2 C less t h a n o v e r m o i s t soil. T h e differences b e t w e e n air
t e m p e r a t u r e s o v e r soil a n d g r a s s m a y h a v e b e e n g r e a t e r t h a n o b s e r v e d h a d the
soil plots b e e n l a r g e r b e c a u s e cold a i r could h a v e d r a i n e d f r o m the s u r r o u n d i n g
g r a s s into the s m a l l e r b a r e areas. T h e i n c r e a s e in a i r t e m p e r a t u r e s a f t e r 03.00 h
(Fig. 3) was c a u s e d by fog.

Leaf temperatures

L e a f t e m p e r a t u r e s e x h i b i t e d b e h a v i o u r s i m i l a r to t h a t of a i r at c o r r e s p o n d -
ing h e i g h t s (Figs. 4 a n d 5). L e a v e s close to the g r o u n d w e r e s i g n i f i c a n t l y colder
t h a n t h o s e at 1000mm a n d g r a d i e n t s in l e a f t e m p e r a t u r e w e r e s t e e p e s t in the
lowest 4 0 m m . S m a l l l e a v e s held h o r i z o n t a l l y at 0 m m a b o v e g r a s s w e r e
t y p i c a l l y 2-3C colder t h a n s i m i l a r leaves at 1000 mm, w h e r e a s leaves at 100 m m
w e r e 1-1.5C colder t h a n a t 1000mm. Clearly, the d u r a t i o n of s u b z e r o tem-
p e r a t u r e s for l e a v e s (and for air, Fig. 1) w a s l o n g e r n e a r the g r o u n d t h a n at
h i g h e r levels (Fig. 4). R a t e s of l e a f cooling n e a r s u n s e t w e r e g r e a t e r close to t h e
s u r f a c e t h a n h i g h e r up a n d l e a v e s cooled m o r e r a p i d l y t h a n t h e a d j a c e n t a i r at
e a c h level.
As for air t e m p e r a t u r e s , f l u c t u a t i o n s in l e a f t e m p e r a t u r e at one h e i g h t w e r e
 127

I'..

::L..

...~-'".....

"..1000
\iO0 "..
. .... T

2 22 "...-":. 0 !". 2 4 6
"~.-~':" ........ "-'i'''}" '?i ..~-.../.<..~.':"-" t (h)
-2L l

-6

Fig. 4. Time series of temperatures of small E. viminalis leaves measured at 0, 40, 100 and 1000 mm
above grass on 2(~ 21 July 1984. Up and down arrows have the same meaning as in Fig. 1.

often observed at all other levels (Fig. 4) and often fluctuations in air and leaf
temperatures had a similar magnitude (Fig. 5). Large and small leaves were
systematically ~ 3 and ~ 2C, respectively, below air temperatures at 100 mm.
Similar differences were observed at 1000mm, but near the ground leaf tem-
peratures were very close to those for air.
Temperature of a leaf (or any other plant organ) on a clear night is
determined by a balance between net radiation (R), fluxes of sensible and latent
heat (H + 2E) through the leaf boundary layer from surrounding air and heat
storage changes in the leaf (Qs). This balance may be written as:
-R = H + )oE + Q~ = pacpkh(l + S/pT~)(T~ - T~)
1 (.
+ ~ | Q(OT~/Ot)dV (2)
,J
v
where kh is the mean boundary layer conductance, S is the slope of the curve
relating air temperature to saturation water vapour pressure over water or ice,
7s is the psychrometric constant for water or ice, p is atmospheric pressure and
T~ and 7'1 are air temperature outside the boundary layer and at the leaf surface,
respectively. Change in energy storage (Q) is given by the last term of eq. 2,
in which A and V are the area and volume of the plant organ, respectively, and
Cl is volumetric heat capacity. This term may be ignored for thin leaves, but is
128

\ ",.

6 ~'
iI '..

4 ~1 "'~""'""":'.

\~ ""'"". air

2 ~ ~/N small "'"'.

,, ,~ ', "I ". ..I.- ...:'j I I I I I

18 20 ,~r22 ".-"0 " .. 2 4. 6
t (h)
V ~,~,~,.--4
V',"
.~ ........../ ~ ,. ~-
-2
\ t ii \1

-4

Fig. 5. Time series of temperatures of air, small (E. viminalis) and large (E. pauciflora) leaves at
100mm above grass on 20-21 July 1984.

important for the energy balance of more massive organs such as stems. Net
radiation at the leaf surface is a function of the absolute temperatures of the
leaf and its surroundings, in our case the sky and the ground. Boundary layer
conductance is a function of leaf size and wind speed, small leaves having a
higher conductance t h a n large leaves at similar wind speeds. Air temperatures
and wind speeds are the same for both leaf sizes at a given height and
inspection of eq. 2 shows t h a t Ta - T1 will be greater for large leaves t han for
small ones for similar values of R. Changes in air t em perat ure will result in
similar changes in leaf t e m per a t ur e since small changes in leaf t em perat ure do
not change R significantly. This is seen in Fig. 5, where fluctuations in air and
leaf temperatures were highly correlated and had similar magnitudes.
For horizontal leaves, the local boundar y layer conductance at the tip or
edge of a leaf is less t ha n the mean kh for both forced and free convection.
Similarly, under conditions of free convection, kh is less for a vertical t han for
a horizontal leaf of the same size. Thus a vertical leaf and a leaf tip will be
closer to air temp er a t ur e t han the centre of a horizontal leaf (eq. 2). An example
of this is shown in Fig. 9.
Ranking of temperatures observed at a given height above grass (Fig. 5) was
large leaves < small leaves < air, and the same order was found for horizontal
leaves displayed above bare soil. For example, at 40 mm above moist soil (Fig.
6), large and small leaves were ~ 2 and IC below air temperature, respectively.
 129

\\ 'l"" ""'

\1,~ ""'...
I~,, "'...

xx . "'".......
\\ ""::air
C mall \
large ~ \ \ ~ _ :'-.... ..
~L
\\ % "".'...... . . . ~ . ..-"/, Y I
0 ( r.. ~ ~ I
18 20 ,, 2 2 ~ },l) I'.... 2 ..9, 6 8
~_ , ~ \
.,./,
'..../J t cht
v x ~ I \ " I
-2 ".' \\ ~ "x'x]///
\ ~ V// \1

\. )
\/
-4

Fig. 6. Time series of temperatures of air, small and large leaves measured at 40 mm above moist
soil on 15- 16 August 1984.

L e a f t e m p e r a t u r e s rose rapidly and c o n v e r g e d with air t e m p e r a t u r e s w h e n fog

developed.
F i g u r e 7 shows the influence of different u n d e r l y i n g surfaces on the tem-
p e r a t u r e of E . v i m i n a l i s leaves placed h o r i z o n t a l l y at 40 mm o v e r grass and over
dry or moist soil. L e a v e s above grass were ~ 1.0 and ~ 1.5C colder t h a n those
above dry and moist soils, respectively, a r e s u l t c o n s i s t e n t with the findings of
L u d l o w and F i s h e r (1976).
H e a t and r a d i a t i o n fluxes at different g r o u n d surfaces d e t e r m i n e the tem-
p e r a t u r e of those surfaces and of the air in c o n t a c t with them. This in t u r n
affects leaf t e m p e r a t u r e s d i r e c t l y by r a d i a t i o n e x c h a n g e with the g r o u n d and
by h e a t c o n d u c t i o n from the air s u r r o u n d i n g the leaf. A less direct influence of
the g r o u n d surface is t h r o u g h its a e r o d y n a m i c roughness, w h i c h affects the
steepness of wind speed and t e m p e r a t u r e profiles n e a r the ground. Oke (1970)
provided several examples of the effect of surface r o u g h n e s s on n o c t u r n a l
t e m p e r a t u r e profiles n e a r the ground.
D i r e c t m e a s u r e m e n t of leaf t e m p e r a t u r e s m a y not be feasible in ecological
studies and often the o n l y d a t a available are air t e m p e r a t u r e s m e a s u r e d in a
s t a n d a r d screen. The above results i n d i c a t e t h a t a simple r e l a t i o n s h i p c a n n o t
be e x p e c t e d b e t w e e n leaf and screen t e m p e r a t u r e s d u r i n g r a d i a t i o n frost.
130

8[
(II

\""

':(i \i"" """

K
E "~. wet soil . ~
0
18 20 \"~.22-.... 0 :" 2 /) 6 8
".. ~'. .[J
~ : " . . . , . !

\ /
-4 ~\ /
dr? soil ,~

Fig. 7. Time series of temperatures of small Leaves measured at 40 mm above grass, dry soil and wet
soil on 1 ~ 1 6 A u g u s t 1984.

! (tIlnl)

O0
la) ]05 {b) 1.0
40
I00
Z~ I 0 0 0
0,4 08
E
/ ,<
0.3 06 e,

-8 6 4 -2
T -T. (C)
i 0
0.2

2 -8
/

6 4
T T (C)
2
0.4

02

Fig. 8. (a) F r e q u e n c y distribution for difference b e t w e e n leaf and screen temperature (T 1 - T~) for
E. viminalis. (b) Cumulative plot for data in Fig. 8a.
 131

However, a useful relation does exist between these temperatures if the results
are presented as frequency distributions (Fig. 8a). Mean screen temperatures
(Ts) were subtracted from the corresponding mean leaf temperatures (T,) to
construct this graph. Data from seven fog-free nights were used and data
selection was limited to periods when Ts < 5C. Distributions of T~ - Ts are
clearly dependent on height above ground. The greatest spread of T~ - Ts
occurred at the grass tips with narrower distributions at 100 and 1000 mm. The
peak of the distributions occurred at progressively higher temperatures with
increased height. Patterns for air temperature and for that of large leaves were
similar to Tl - Ts for small leaves.
Cumulative frequencies derived from the same data as for Fig. 8a are
presented in Fig. 8b since these are suitable for predictive purposes. At 40 mm
for example, TI was > 3.6C below Ts for 50% of the time. No leaves were > 9C
below screen temperature and TI was greater than Ts for only 5% of the time
whenever screen temperatures were < 5C.
The frequency distributions of Fig. 8 apply to the particular case of isolated
leaves displayed horizontally over a grass sward and different distributions
should be expected for other arrangements. Screen (and grass-minimum)

Horizontal
8OO

large small r

Vertical leaves /
0 0 large 500
~- ~ small Z

(ram)

400

200

~ 0
-5 -4 -3 -2 I 0
T (C)
Fig. 9. Typical temperature profiles for air, small leaves and large horizontal leaves on 2(~21 July
1984 when T, was at its lowest temperature for the night. The effect of vertical orientation on leaf
temperature is shown.
132

t e m p e r a t u r e s p r o v i d e o n l y a guide to the t e m p e r a t u r e s a t t a i n e d by p l a n t
organs. A p p a r e n t l y m i n o r differences in the m i c r o h a b i t a t or m o r p h o l o g y of
p l a n t p a r t s (e.g., h e i g h t a b o v e ground, size, o r i e n t a t i o n , g r o u n d s u r f a c e type)
m a y r e s u l t in s i g n i f i c a n t v a r i a t i o n s in t h e i r t e m p e r a t u r e s d u r i n g n i g h t s of
r a d i a t i o n frost. I n v e s t i g a t o r s a t t e m p t i n g to c o r r e l a t e d a m a g e or d e a t h of p l a n t
o r g a n s w i t h t e m p e r a t u r e s m e a s u r e d e l s e w h e r e s h o u l d be a w a r e of t h e s e f a c t o r s
w h e n i n t e r p r e t i n g t h e i r results.
R e p r e s e n t a t i v e profiles for a i r a n d l e a f t e m p e r a t u r e s s u m m a r i z i n g the m a i n
o b s e r v a t i o n s of this s t u d y are p r e s e n t e d in Fig. 9. T h e m a i n points are: (1) at
a n y g i v e n height, the c e n t r e s of l a r g e r l e a v e s w e r e cooler t h a n t h o s e of t h e
s m a l l e r l e a v e s a n d b o t h w e r e colder t h a n air; (2) l e a f a n d air t e m p e r a t u r e s w e r e
lowest n e a r the g r o u n d a n d i n c r e a s e d w i t h height; (3) t e m p e r a t u r e g r a d i e n t s
w e r e v e r y steep n e a r the g r o u n d ( ~ 40C m 1 for air, ~ 20C m 1 for small leaves
and ~ 5C m 1 for l a r g e l e a v e s b e t w e e n 0 a n d 100mm); (4) l e a f a n d air tern-
p e r a t u r e s t e n d e d to c o n v e r g e at the surface, a n d (5) l a r g e leaves held v e r t i c a l l y
w e r e ~ IC closer to a i r t e m p e r a t u r e t h a n h o r i z o n t a l l e a v e s at the s a m e height.
A lesser effect of o r i e n t a t i o n w a s f o u n d for small leaves. A l t h o u g h n o t s h o w n
in Fig. 9, the t y p e of u n d e r l y i n g s u r f a c e also influences the t e m p e r a t u r e of the
leaves a b o v e t h e m (Fig. 7).
This p a p e r h a s identified s o m e of t h e m a j o r f a c t o r s a f f e c t i n g t e m p e r a t u r e s
a t t a i n e d by i s o l a t e d leaves of b r o a d - l e a v e d seedlings. T h e o r y w h i c h e x p l a i n s
q u a n t i t a t i v e l y leaf-to-air t e m p e r a t u r e differences and the t e n d e n c y of l e a f and
air t e m p e r a t u r e s to c o n v e r g e at the g r o u n d is p r e s e n t e d in a c o m p a n i o n p a p e r
(Leuning, 1987).

CONCLUSIONS

L e a f t e m p e r a t u r e s d u r i n g periods of r a d i a t i o n frost are d e t e r m i n e d by a

c o m b i n a t i o n of e n v i r o n m e n t a l a n d p l a n t m o r p h o l o g i c a l f a c t o r s t h a t affect the
b a l a n c e b e t w e e n n e t loss of e n e r g y f r o m t h e l e a f by r a d i a t i o n to the s k y and
g a i n by h e a t c o n d u c t i o n f r o m t h e s u r r o u n d i n g a i r a n d by r a d i a t i o n f r o m a n y
w a r m e r n e i g h b o u r i n g t e r r e s t r i a l surfaces.

REFERENCES

Andre, J.C. and Mahrt, L., 1982. The nocturnal surface inversion and influence of clear-air
radiative cooling. J. Atmos. Sci., 39: 864~878.
Cannell, M.G.R., 1984. Spring frost damage on young Picea sitchensis. I. Occurrence of damaging
frosts in Scotland compared with Western North America. Forestry, 57: 159-175.
Coantic, M. and Seguin, B., 1971. On the interaction of turbulent and radiative transfers in the
surface layer. Boundary-Layer Meteorol., 1: 24~263.
Funk, J.P., 1960. Measured radiative flux divergence near the ground at night. Q. J. R. Meteorol.
Soc., 86:382 389.
Lake, J.V., 1956. The temperature profile above bare soil on clear nights. Q. J. R. Meteorol. Soc.,
82:187 197.
Leuning, R., 1987. Leaf temperatures during radiation frost. Part II. A steady-state theory. Agric.
For. Meteorol., 42: 135-155.
 133

Levitt, J., 1980. Responses of Plants to Enviromental Stresses. Vol. 1. Chilling, Freezing, and High
Temperature Stresses. Academic Press, New York.
Ludlow, M.M. and Fisher, M.J., 1976. Influence of soil surface litter on frost damage. Aust. Inst.
Agric. Sci. J., 42: 134-136.
Nunez, M. and Sander, D., 1982. Protection from cold stress in a Eucalyptus shelterwood. J.
Climatol., 2: 141-146.
Ogawa, Y., Diosey, P.G., Uehara, K. and Ueda, H., 1985. Wind tunnel observation of flow and
diffusion under stable stratification. Atmos. Environ., 19: 6~74.
Oke, T.R., 1970. The temperature profile near the ground on calm clear nights. Q. J. R. Meteorol.
Soc., 96:14 23.
Steponkus, P.L., 1978. Cold hardiness and freezing injury of agronomic crops. Adv. Agron., 30:
51 98.