Journal of Experimental Botany, Vol. 50, No. 336, pp.
1199–1206, July 1999
Effects of water stress on photosystem II photochemistry
and its thermostability in wheat plants
Congming Lu and Jianhua Zhang1
Department of Biology, Hong Kong Baptist University, Kowloon Tong, Hong Kong
Received 23 October 1998; Accepted 22 February 1999
Abstract
Modulated chlorophyll fluorescence, rapid fluores-
cence induction kinetics and the polyphasic fluores-
cence transients (OJIP) were used to evaluate PSII
photochemistry in wheat plants exposed to water
stress and/or heat stress (25–45 °C). Water stress
showed no effects on the maximal quantum yield of
PSII photochemistry (F /F ), the rapid fluorescence
v m
induction kinetics, and the polyphasic fluorescence
transients in dark-adapted leaves, indicating that
water stress had no effects on the primary photochem-
istry of PSII. However, in light-adapted leaves, water
stress reduced the efficiency of excitation energycap-
ture by open PSII reaction centres (F∞ /F∞ ) and
v m
the quantum yield of PSII electron transport (w ),
PSII
increased the non-photochemical quenching (q ) and
N
showed no effects on the photochemical quenching
(q ). This suggests that water stress modified the PSII
P
photochemistry in the light-adapted leaves and such
modifications may be a mechanism to down-regulate
the photosynthetic electron transport to match a
decreased CO assimilation. In addition, water stress
2 2
also modified the responses of PSII to heat stress.
When temperature was above 35 °C, thermostability of
PSII was strongly enhanced in water-stressed leaves,
which was reflected in a less decrease in F /F , q ,
v m P
F∞ /F∞ , and w in water-stressed leaves than in well-
v m PSII
watered leaves. There were no significant variations
in the above fluorescence parameters between moder-
ately and severely water-stressed plants, indicating
that the moderate water stress treatment caused the
same effects on thermostability of PSII as the severe
treatment. It was found that increased thermostability
of PSII may be associated with an improvement of
resistance of the O -evolving complex and the reaction
2
centres in water-stressed plants to high temperature.
1 To whom correspondence should be addressed. Fax: +852 2336 1400. E-mail: jzhang@hkbu.edu.hk
© Oxford University Press 1999
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Key words: Chlorophyll fluorescence, heat stress, photo-
system II photochemistry, water stress, wheat (Tritium
aestivum L.).
Introduction
Water stress is one of the most important environmental
factors inhibiting photosynthesis (Bradford and Hsiao,
1982). Many studies have shown that the decreased
photosynthesis under water stress can be associated with
the perturbations of the biochemical processes (Graan
and Boyer, 1990; Lauer and Boyer, 1992). In particular,
PSII has been shown to be very sensitive to water stress.
Several in vivo studies demonstrated that water stress
resulted in damage to the oxygen-evolving complex of
PSII (Canaani et al., 1986; Toivonen and Vidaver, 1988)
and to the PSII reaction centres (Havaux et al., 1987; He
et al., 1995).
On the other hand, other studies have shown that the
inhibited CO -dependent O evolution and net CO assim-
2 2 2
ilation induced by water stress can be recovered by high
external CO concentration ( Frederik et al., 1990; Cornic,
1994), implying that the perturbations of the biochemical
processes are not responsible for the inhibited CO assim-
2
ilation, and stomata instead may play a dominant role in
the decreased CO assimilation under water stress. The
2
conclusion can be further supported by the fact that PSII
photochemistry is hardly affected by water stress (Cornic
and Briantais, 1991; Cornic, 1994; Liang et al., 1997).
Thus, it is still a matter of uncertainty how water stress
affects PSII photochemistry.
Under natural conditions, multiple environmental
stresses co-occur frequently. It has been reported that the
responses of plants to several simultaneous stresses are
usually not predictable by single-factor analysis and a
combination of different environmental stress factors can
1200 Lu and Zhang
result in intensification, overlapping or antagonistic effects
(Osmond et al., 1986). The effects of interaction of
drought and high light on PSII have been investigated
widely, showing that drought predisposes photoinhibitory
damage to PSII (Masojidek et al., 1991; Giardi et al.,
1996). However, much less attention has been paid to the
responses of PSII to a combination of water stress and
heat stress. It is obviously important to know the mechan-
isms of the interaction of these two stresses on PSII
functions from the view of an ecophysiological aspect,
since water stress is often combined with heat stress, in
particular in arid and semi-arid regions.
In this study, the effects of water stress on PSII
photochemistry and thermostability of PSII in wheat
plants by analyses of fluorescence quenching, the rapid
fluorescence induction kinetics (i.e. Kautsky curve), and
the polyphasic rise of fluorescence transients were exam-
ined ( Krause and Weis, 1991; Schreiber et al., 1994;
Govindjee, 1995; Strasser et al., 1995; Strasser, 1997). It
is demonstrated that, although water stress showed no
effect on the primary photochemistry of PSII, it increased
thermostability of PSII. It is further demonstrated that
such increased thermostability may be associated with an
improvement on resistance of the O -evolving complex
2
and the reaction centres to high temperature.
Materials and methods
Plant material and stress treatments
Wheat (Tritium aestivum L. cv. Shannong 229) seedlings were
grown in plastic pots (14 cm in diameter and 13 cm in height)
in a growth chamber. The pots were filled with soil which was
composed of loam, peat, and coarse sand in 75352 volume
ratio and added with NPK (15515515, by vol.) complete
fertilizer. Light irradiance at the top of plants was c.
180 mmol m−2 s−1 as provided by high pressure sodium lamps
(SON-T AGRO, Philips, Belgium) with a photoperiod of
16/8 h. Air temperature during day and night was kept at
20±1 °C and relative humidity at 80±3%. All plants were
watered daily.
After 4 weeks, the seedlings were used for the water stress
treatment, which was developed gradually by withholding
water. After a period of 10 d, leaf water potential reached
−2 MPa, measured with a pressure chamber (Model 3000,
Soil Moisture Equipment Co., USA). The youngest and fully
expanded leaves were used for the measurements.
Heat stress was applied on leaf discs (2 cm). The leaf disc
was directly placed into the smooth bottom of a small hole
(5.5 cm in height×3 cm in diameter) in a block of brass, of
which the temperature was regulated by circulation of water
from a thermostated water bath. At the same time, the up-face
of the leaf dics was pressed directly by a block of glass so that
the water evaporation of the leaf could be prevented and the
heat equilibrium between the leaf disc and the brass block
could be reached immediately. The hole of the brass was also
covered during treatment. The leaf discs were exposed to
different elevated temperatures in the dark for 15 min and then
their fluorescence characteristics were measured at 25 °C before
and after temperature treatment. No difference was observed in
the rate of temperature increase between drought-stressed and
well-watered leaves. Also, no changes were observed in various
fluorescence parameters during the measurements of fluores-
cence after dark-adaptation of heat-stressed leaves.
Analysis of gas exchange
Gas exchange analysis was made using an open system (Ciras-1,
PP system, UK ). Net CO assimilation rate was determined at
2
a CO concentration of 360 cm3 m−3, 80% relative humidity,
2
and 180 mmol m−2 s−1 light intensity. Leaf stomatal conduct-
ance (G ) was measured under the same conditions with a
s
steady-state porometer (Li-1600, Li-Cor, Lincoln, NE, USA).
Measurements of chlorophyll fluorescence
Chlorophyll fluorescence was measured at room temperature
with a portable fluorometer (PAM-2000, Walz, Germany). The
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fluorometer was connected to a leaf-clip holder (2030-B, Walz)
with a tri-furcated fibre optic (2010-F, Walz) and to a computer
with data acquisition software (DA-2000, Walz). The experi-
mental protocol of Genty et al. (1989) was basically followed.
The minimal fluorescence level (F ) with all PSII reaction
0
centres open was measured by the measuring modulated light
which was sufficiently low (<0.1 mmol m−2 s−1) not to induce
any significant variable fluorescence. The maximal fluorescence
level (F ) with all PSII reaction centres closed was determined
m
by a 0.8 s saturating pulse at 8000 mmol m−2 s−1 in dark-
adapted leaves. The leaf disc was then continuously illuminated
with white actinic light at an intensity of 180 mmol m−2 s−1
which was equivalent to growth PPFD of wheat seedlings in
the growth chamber. The steady-state value of fluorescence (F )
s
was thereafter recorded and a second saturating pulse at
8000 mmol m−2 s−1 was imposed to determine the maximal
fluorescence level in light-adapted leaves (F∞ ). The actinic light
m
was removed and the minimal fluorescence level in the light-
adapted state (F∞ ) was determined by illuminating the leaf disc
0
with 3 s far-red light. All measurements of F and F∞ were
0 0
performed with the measuring beam set to a frequency of
600 Hz, whereas all measurements of F and F∞ were performed
m m
with the measuring beam automatically switching to 20 kHz
during the saturating flash.
By using fluorescence parameters determined in both light-
and dark-adapted leaves, the following calculations were made
(1) the maximal quantum yield of PSII photochemistry,
F /F , (2) the photochemical quenching coefficient, q =
v m P
(F∞ −F )/(F∞ −F∞ ), and the non-photochemical quenching
m s m 0
coefficient, q =1−(F∞ −F∞ /(F −F ), (3) the efficiency of
N m 0 m 0
excitation capture by open PSII reaction centres, F∞ /F∞ , and
v m
the actual quantum yield of PSII electron transport, w .
PSII
Fluorescence nomenclature was according to van Kooten and
Snel (1990).
The rapid fluorescence induction kinetics ( Kautsky curve)
were measured in dark-adapted leaves suddenly illuminated
with moderate red light (peak at 655 nm) of 45 mmol m−2 s−1
at a sampling rate of 1000 ms point−1. In order to avoid an
incomplete reoxidation of the plastoquinone pool in the dark
which could result in an increase in fluorescence level at phase
I, the dark-adapted leaves were illuminated with 3 s far-red
light prior to measurement of the fluorescence induction kinetics.
The polyphasic rise of fluorescence transients (OJIP) was
measured by a Plant Efficiency Analyser (PEA, Hansatech
Instruments Ltd., King’s Lynn, UK ) according to Strasser
et al. (1995). The transients were induced by red light of
3000 mmol m−2 s−1 provided by an array of six light-emitting
diodes (peak 650 nm), which focused on the sample surface to
give homogeneous illumination over the exposed area of the

sample (4 mm in diameter). The fluorescence signals were
recorded within a time scan from 10 ms to 1 s with a data
acquisition rate of 100 000 points s−1 for the first 2 ms and of
1000 points s−1 after 2 ms. The fluorescence signal at 40 ms was
considered as a true F since the fluorescence yield at this time
0
was shown to be independent of light intensity.
All samples were dark-adapted for 30 min prior to fluores-
cence measurements.
Results
Effects of water stress on water status, net CO assimilation
2 of chlorophyll fluorescence from the minimal level (O) to
and stomatal conductance
Water stress was induced gradually in wheat plants by
withholding water. Two levels of water stress were exam-
PSII thermostability in water-stressed plants 1201
Although F /F is often used to assess the maximal
v m
quantum yield of PSII photochemistry and shows no
change during water stress, it gives no direct information
on the heterogeneity of PSII reaction centres. In order to
examine further whether water stress induces the changes
in the heterogeneity of PSII reaction centres, the rapid
fluorescence induction kinetics was determined.
When a dark-adapted leaf was illuminated with red
light of moderate intensity (40 mmol m−2 s−1), a typical
Kautsky curve was observed, which displayed a rapid rise
an intermediate level (I ) followed by a very fast rise to
the maximum level (P) (the inset in Fig. 1). The O-I
phase has been attributed to Q reduction in the Q -non-
A B

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ined, moderate and severe water stress, which were
reached c. 3–4 d and 10 d, respectively, after withholding
water. Plants wilted if further water stress was imposed
after 10 d. Water stress decreased the relative water con-
tent of leaves ( RWC ) and the leaf water potential (y ).
w
RWC decreased from 92.1% in well-watered plants to
88.3% and 80.5% in moderately and severely water-
stressed plants, respectively. Accordingly, y decreased
w
from −0.56 in well-watered plants to −1.12 and −1.95
MP in moderately and severely water-stressed plants,
a B
respectively ( Table 1).
The rate of net CO assimilation decreased by c. 52%
2
and 89% in moderately and severely water-stressed plants,
respectively. Leaf stomatal conductance also decreased
significantly as water stress developed ( Table 1).
Effects of water stress on PSII photochemistry
No changes were observed in the maximal quantum yield
of PSII photochemistry (F /F ) measured in dark-adapted
v m
leaves in either moderate or severely stressed plants
( Table 2).
reducing PSII reaction centres, in which the electron
transfer from Q− to Q is inhibited and phase I-P reflects
A B
the accumulation of Q− in the active PSII reaction centres
A
with efficient electron transfer to the plastoquinone pool
(Chylla and Whitmarsh, 1989; Cao and Govindjee, 1990).
The ratio (F −F )/(F −F ) can thus be considered a
i 0 p 0
measure of the percentage of those Q -non-reducing PSII
B
reaction centres. Table 2 and Fig. 1 show that water stress
induced no changes in this ratio, suggesting that water
stress exerted no effect on the proportion of Q -non-
reducing PSII reaction centres.
Recently, it has been demonstrated that when a dark-
adapted leaf is illuminated suddenly with high intensity
actinic light (3000 mmol m−2 s−1), the fluorescence transi-
ent shows a polyphasic rise, including phases O, J, I, and
P (Strasser et al., 1995). The initial Chl fluorescence at
O level reflects the minimal fluorescence yield when all
molecules of Q are in the oxidized state. The P level
A
corresponds to the state in which all molecules of Q are
A
in the reduced state. Steps J and I occur at about 2 ms
and 30 ms, respectively, between the commonly observed

Table 1. Effects of water stress on water potential (Y ), the relative water content of leaves (RWC), stomatal conductance (G ),
w s
and net CO assimilation rate (A) in the leaves of wheat plants; values are means±SE of 4–8 replicates
2
Y RWC G A
w s
(MPa) (%) (mmol m−2 s−1) (mmol m−2 s−1)

Well-watered 0.56±0.07 92.1±4.3 220.2±8.9 15.5±0.3

Moderately stressed 1.12±0.12 88.3±5.2 105.9±12 7.5±0.3
Severely stressed 1.95±0.09 80.5±8.0 50.7±6.5 1.7±0.06

Table 2. Effects of water stress on the maximal quantum yield of photochemistry of PSII (F /F ) and the ratio (F −F )/(F −F )
v m i 0 p 0
determined from the rapid fluorescence induction kinetics in dark-adapted leaves, photochemical (q ) and non-photchemical (q )
P N
quenching, the efficiency of excitation capture by open PSII reaction centres (F∞ /F∞ ) and the actual quantum yield of PSII electron
v m
transport (W ) in light-adapted leaves, which were illuminated by white actinic light at an intensity of 180 mmol m−2 s−1: values are
PSII
means±SE of four replicates

F /F (F −F )/(F −F ) q q F∞ /F∞ W
v m i 0 p 0 P N v m PSII
Well-watered 0.821±0.007 0.232±0.012 0.781±0.008 0.321±0.005 0.752±0.008 0.601±0.009
Moderately stressed 0.824±0.008 0.221±0.020 0.790±0.005 0.367±0.003 0.676±0.003 0.545±0.005
Severely stressed 0.828±0.006 0.226±0.017 0.785±0.008 0.448±0.003 0.610±0.006 0.501±0.006
1202 Lu and Zhang

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Fig. 1. Changes in the proportion of Q -non-reducing PSII reaction
B
centres, expressed as (F −F )/(F −F ), in well-watered ($) and severely
i 0 p 0
water-stressed (&) leaves exposed to elevated temperatures in the dark
for 15 min. Each value is mean±SE of four samples. The inset: the
rapid fluorescence induction kinetics (O-I-P) in dark-adapted well-
watered leaves suddenly illuminated with moderate red light
(40 mmol m−2 s−1, peak at 655 nm).

steps O and P. The rise from phase O to phase J results

from the reduction of Q to Q− and is associated with
A A
the primary photochemical reactions of PSII. The inter-
mediate step I and the final step P reflect the existence of
fast and slow reducing PQ centres as well as to the
different redox states of PSII reaction centres (Strasser
et al., 1995). Thus, the polyphasic rise of fluorescence
transients can give more information on PSII photo-
chemistry.
The polyphasic Chl a fluorescence transients in wheat
plants during water stress were followed. The polyphasic
rise in Chl a fluorescence transients in well-watered plants
demonstrated a typical polyphasic rise, including phases
O, J, I, P, described in details by Strasser et al. (1995).
It was observed that water stress had no significant effect
on these phases in the polyphasic rise of fluorescence
transients ( Fig. 2A, curve a, b).
The above results show that water stress had no effects
on the primary photochemistry of PSII determined in
dark-adapted leaves. It was further investigated whether
water stress induces modifications in PSII photochemistry
in light-adapted leaves. The changes in fluorescence char-
acteristics in light-adapted leaves were examined by the
use of pulse-modulated fluorescence.
Table 2 shows that water stress decreased the efficiency
of excitation energy capture by open PSII reaction centres
(F∞ /F∞ ) and the quantum yield of PSII electron transport
v m
(w ), increased the non-photochemical quenching
PSII
Fig. 2. (A) Effects of high temperature on the polyphasic chlorophyll
fluorescence transients in well-watered and severely water-stressed
leaves. The curves are (a) well-watered leaves, (b) severely water-
stressed leaves, (c) severely water-stressed leaves exposed to 45 °C for
15 min, and (d ) well-watered leaves exposed to 45 °C for 15 min. (B)
Changes in the ratio F /F (i.e. the ratio of fluorescence level at 300 ms
K J
to the fluorescence level at 2 ms) in well-watered ($) and severely
water-stressed (&) leaves after exposed to elevated temperatures in the
dark for 15 min. Each value is mean±SE of four samples.
coefficient (q ). However, it had no effects on the photo-
N
chemical quenching coefficient (q ).
P
Effects of water stress on thermostability of PSII
The effects of water stress on thermostability of PSII were
investigated. First, the effects of the degree of water stress
on thermostability of PSII were examined. No significant
difference was observed in the responses of PSII photo-
chemistry (the first parameters in dark- and light-adapted
leaves, such as F /F , F∞ /F∞ , w , q , and q , data not
v m v m PSII P N
shown) to heat stress between moderate and severely
water-stressed plants, suggesting that the moderate water
stress treatment caused the same effects on thermostability
of PSII as the severe treatment. Therefore, only the results
in severely water-stressed plants are presented here.
The responses of the maximum quantum yield of PSII

photochemistry (F /F ), the minimal fluorescence (F )
v m 0
and the maximum fluorescence (F ) in dark-adapted
m v m
leaves to elevated temperatures are shown in Fig. 3. When
temperature was increased to 40 °C, F /F began to
v m
decrease significantly in both well-watered and water-
stressed plants. More importantly, water-stressed plants
exhibited higher values of F /F than well-watered plants
v m
under high temperatures. For example, at 45 °C, F /F
v m
decreased by 38% in well-watered plants but only 18% in
water-stressed plants, indicating that water stress induced
an increase in resistance of PSII to heat stress ( Fig. 3A).
A larger increase in F and a greater decrease in F were
0 m
observed in well-watered leaves than in water-stressed
leaves ( Fig. 3B, C ).
It has been suggested that PSII reaction centre is one
of the damage sites induced by heat stress (Havaux,
Fig. 3. Changes in the maximum quantum yield of PSII photochemistry
(F /F , A), the minimum fluorescence yield (F , B), and the maximum
v m 0
fluorescence yield (F , C ) in well-watered ($) and severely water-
m These results show that no significant differences in the
stressed (&) leaves exposed to elevated temperatures in the dark for
15 min. Each value is mean±SE of four measurements.
PSII thermostability in water-stressed plants 1203
1993). Therefore, this study investigated whether the
greater decrease in F /F in well-watered plants during
heat stress was associated with a larger increase in the
proportion of Q -non-reducing PSII reaction centres,
B
expressed as the ratio (F −F )/(F −F ), which has been
i 0 p 0
shown to be sensitive to heat stress ( Klinkovsky and
Naus, 1994). Figure 1 shows that indeed a larger increase
in this ratio was observed in well-watered plants than in
water-stressed plants, suggesting that water stress also
increased thermostability of PSII reaction centres.
It has been shown that the O -evolving complex (OEC )
2
is most susceptible in PSII apparatus to heat stress (Berry
and Björkman, 1980). This study further examined
whether the greater decrease in F /F in well-watered
v m
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plants during heat stress was associated with the greater
inhibition of the OEC activity.
As shown in Fig. 2, well-watered plants show a typical
polyphasic rise of fluorescence transients, including phases
O, J, I, P. It has been shown that heat stress can induce
a rapid rise in the polyphasic fluorescence transients. This
phase, occurring at around 300 ms, has been labelled as
K, and is the fastest phase observed in the OJIP transient
which, in consequence, becomes an OKJIP transient
(Guissé et al., 1995; Srivastava and Strasser, 1996;
Srivastava et al., 1997). It has also been shown that phase
K is caused by an inhibition of electron donor to the
secondary electron donor of PSII, Y , which is due to a
Z
damaged OEC, phase K can therefore be used as a specific
indicator of damage to the OEC (Guissé et al., 1995;
Strasser, 1997).
Figure 2A shows that when exposed to 45 °C for 15 min,
a very clear K step appeared in well-watered plants. But,
this heat-induced K step did not get pronounced in water-
stressed plants. The changes in the amplitude in the K
step, expressed as the ratio F /F , was shown in Fig. 2B.
K J
A higher ratio F /F was observed in well-watered plants
K J
than in water-stressed plants during 35–45 °C, indicating
that water stress can increase resistance of the OEC to
heat stress.
Further experiments were carried out to examine the
responses of PSII photochemistry in light-adapted leaves,
i.e. under steady-state photosynthesis, to high temper-
atures. Figure 4 shows that a greater decrease in w ,
PSII
F∞ /F∞ , and q was observed in well-watered plants than
v m P
in water-stressed plants in the range 35–45 °C, indicating
that PSII in water-stressed plants also shows better ther-
mostability in light-adapted leaves.
Discussion
Water stress has no effect on the primary photochemistry of
PSII
maximal quantum yield of PSII photochemistry (F /F ),
v m
1204 Lu and Zhang
Fig. 4. Changes in the quantum yield of PSII electron transport (w ,
A), the efficiency of excitation capture by open PSII reaction centres
(F∞ /F∞ , B), and the photochemical quenching coefficient (q , C ) in well-
v m
watered ($) and severely water-stressed (&) leaves exposed to elevated
temperatures in the dark for 15 min. Each value is mean±SE of
four samples.
and in the rapid fluorescence induction kinetics and the
polyphasic rise of fluorescence transients were observed
between well-watered and water-stressed plants, indicat-
ing that water stress had no effect on the primary photo-
chemistry of PSII (Table 2; Figs 1, 2). However, these
results show that water stress induced a decrease in the
quantum yield of PSII electron transport (w ) ( Table 2),
PSII
suggesting that water stress resulted in modifications
in PSII photochemistry in light-adapted leaves. The
decreased w was a result of the decrease in the efficiency
PSII
of excitation energy capture by open PSII reaction centres
(F∞ /F∞ ) because of no change in photochemical quenching
v m
(q ) (w =F∞ /F∞ ×q , Genty et al., 1989) ( Table 2).
P PSII v m P
A decrease in F∞ /F∞ may reflect light-induced non-
v m
photochemical quenching (Baker, 1991). An increase in
q was indeed observed in water-stressed plants. This
N
increased q would dissipate some excitation energy at
N
the expense of photochemical utilization (Brestic et al.,
1995). Thus, a high q in water-stressed leaves may be a
N
mechanism to down-regulate photosynthetic electron
transport so that production of ATP and NADPH would
match with the decreased CO assimilation due to the
2
closure of stomata ( Table 1).
Water stress increases thermostability of PSII
The results in this study demonstrate the existence of an
antagonism between water stress and heat stress in wheat
plants, with water stress enhancing the resistance of PSII
to heat stress. It was found that increased resistance of
PSII to heat stress in water-stressed leaves was exhibited
not only in the resistance of the maximal quantum
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efficiency of PSII photochemistry in the dark-adapted
state (i.e. F /F ) ( Fig. 3A) as previously observed by
v m
Havaux (1992) and Epron (1997), but also in the resist-
ance of PSII photochemistry in the light-adapted state,
i.e. under steady-state photosynthesis, such as F∞ /F∞ , q ,
v m P
and (PSII, to high temperatures ( Fig. 4).
These results show that such an increased resistance of
PSII to heat stress in water-stressed leaves was associated
with an improvement of the resistance of PSII reaction
centres to high temperatures, as indicated by a smaller
increase in the proportion of Q -non-reducing PSII reac-
B
tion centres during heat stress in water-stressed leaves
than in well-watered plants (Fig. 1). A smaller decrease
in photochemical quenching (q ) also suggests increased
P
thermostability of PSII reaction centres in water-stressed
plants since q can represent the fraction of open PSII
P
reaction centres (Genty et al., 1989; van Kooten and Snel,
1990) (Fig. 4C ). In addition, these results demonstrate
PSII
that increased thermostability of PSII in water-stressed
P plants was also associated with an improvement in ther-
mostability of the O -evolving complex (OEC ), as shown
2
by a less pronounced phase K in the polyphasic fluores-
cence transients in water-stressed plants than in well-
watered plants (Fig. 2).
How water stress can protect PSII against heat damage
is not clear. One possibility is that increased thermostabil-
ity of PSII may be associated with the accumulation of
some soluble compounds in water-stressed leaves. It has
been shown that thermoresistance of the thylakoids to
high temperature stress is enhanced, when the chloroplasts
are incubated in the presence of some soluble compounds,
such as sugars and proteins ( Krause and Santarius, 1975;
Santarius, 1975). It is interesting that these soluble
compounds indeed accumulate in water-stressed leaves
(Seemann et al., 1986; Sachs and Ho, 1986).
Water stress usually results in a decrease in leaf transpir-
ation because of the stomatal closure which commonly
occurs under water stress. Such a decrease in leaf transpir-
ation results in an increase in leaf temperature. However,
it has been shown that leaf photosynthesis is thought to
be the main target of high temperatures and PSII within

the photosynthetic apparatus appears to be most heat
sensitive (Berry and Björkman, 1980). Increased thermo-
stability of PSII in water-stressed leaves may thus help to
improve the resistance of the photosynthetic metabolism
to high temperature and thus may increase the resistance
of the whole plant to high temperature.
Acknowledgements
We are grateful to Hong Kong Baptist University (FRG grant)
and the Croucher Foundation for their financial support to
this study.
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