Tree Physiology 18, 607--614
1998 Heron Publishing----Victoria, Canada
Photosynthesis and photoprotection inQuercus ilex resprouts after fire
I. FLECK,1 K. P. HOGAN,2 L. LLORENS,1 A. ABADA3 and X. ARANDA1
1
Dept. Biologia Vegetal, Unitat Fisiologia Vegetal, Facultat Biologia, Univ. Barcelona, Diagonal 645, 08028 Barcelona, Spain
2
Manaaki Whenua Landcare Research, P.O. Box 69, Lincoln 8152, New Zealand
3
Dept. Nutricin Vegetal, Estacin Experimental de Aula Dei, CSIC, Apdo. 202, 50080 Zaragoza, Spain
Received July 24, 1997
Summary Plants that resprout after fires often have higher
rates of photosynthesis than before a fire. To elucidate the
mechanism of this response, we studied gas exchange and
chlorophyll fluorescence in Quercus ilex L. plants growing on
control (unburned) sites and on sites that had been burned the
preceding summer. In early July, photosynthetic rates and
stomatal conductance were similar in plants on unburned and
burned plots, and in young and old foliage within unburned
plots. At this time, photochemical efficiency of photosystem II
(PSII), nonphotochemical quenching of chlorophyll fluores-
cence (NPQ), and the de-epoxidation of violaxanthin to zeax-
anthin were also similar among leaves of different ages and
treatments. In late July, photosynthetic rates and stomatal
conductances were much greater in resprouts on the burned
areas than in unburned plants. From early to late July, unburned
plants showed an increase in NPQ and the de-epoxidation of
violaxanthin to zeaxanthin, indicating increased photoprotec-
tion as a result of enhanced nonradiative dissipation of excess
light energy. Plants on the burned plots did not show these
changes. Leaves of all ages and treatments showed no substan-
tial reduction in potential quantum yield of PSII (Fv /Fm) at
midday or predawn, indicating that there was little or no
photoinhibition. Leaf nitrogen and soluble protein contents
varied with leaf age during July, but did not vary between
treatments. We conclude that the primary effect of burning is
an increase in water availability to resprouting plants that
eliminates the need for photoprotection, at least in the short
term. The decrease in photosynthetic rates of unburned leaves
in late July was the result of reduced stomatal conductance. We
suggest that lowered stomatal conductance is the primary lim-
iting factor in Q. ilex leaves, governing the regulation of
carboxylation activity and energy dissipation processes.
Keywords: chlorophyll fluorescence, gas exchange, Holm oak,
photoinhibition, photosystem II, zeaxanthin.
Introduction
Holm oaks (Quercus ilex L.) are frequently subjected to stress-
ful environmental conditions including drought, high tempera-
tures and fire. A knowledge of the physiology of resprouting
vegetation after fire may help to elucidate mechanisms under-
lying this species adaptive capacity. Such knowledge may also
be important in the future because changes in climate are likely
to induce changes in factors that control the fire regime (Davis
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and Michaelsen 1995).
Resprouting from lignotubers (site of latent buds and stored
carbohydrates) is a common mechanism of regeneration in
Mediterranean shrub ecosystems after a disturbance, and per-
mits vegetation to fill available space rapidly (Trabaud and
Mthy 1988). Enhancement of gas exchange and growth dur-
ing the first growing season after fire has been reported in
chaparral species (e.g., Schlesinger and Gill 1980, Oechel and
Hastings 1983). As a result of reduced plant competition after
a fire, resprouts have more available solar radiation (Hulbert
1988) and nutrients, especially nitrogen (Hastings et al. 1989).
Moreover, reductions in leaf area index (LAI) in the first years
after fire (Rundel and Parsons 1979) result in increased water
availability to the growing resprouts (De Souza et al. 1986).
Recovery after fire often occurs during the summer period
when soil and atmospheric water deficits are high resulting in
a midday depression of photosynthesis (Tenhunen et al. 1987).
In addition, Mediterranean species may be predisposed to
photoinhibition (Demmig-Adams et al. 1989, Quick et al.
1992) because absorption of light may be in excess of that
required for carbon assimilation. It has been suggested that the
dissipation of excitation energy as heat by zeaxanthin, formed
in the xanthophyll cycle by de-epoxidation of violaxanthin,
protects the photosynthetic apparatus from photodamage by
diverting a proportion of the light energy away from the reac-
tion centers (Demmig-Adams and Adams 1992, Horton and
Ruban 1992). Changes in zeaxanthin content are closely cor-
related with changes in non-photochemical quenching of chlo-
rophyll fluorescence (NPQ) (Bilger and Bjrkman 1990,
Demmig-Adams and Adams 1993, Gilmore and Yamamoto
1993). However, Johnson et al. (1993) found that the capacity
for nonradiative energy dissipation, as indicated by NPQ, did
not correlate well with the ecological status of several plant
species.
Previous studies on Mediterranean species that had re-
sprouted after fire showed that, under midday conditions in
summer, photosynthetic rates and stomatal conductance were
higher in resprouts than in plants in unburned areas (Fleck et
al. 1995, 1996a, 1996b). The aim of this study was to assess
whether Quercus ilex leaves of plants in unburned plots dif-
608
fered in susceptibility to photoinhibition and in photoprotec-
tive mechanisms at midday from resprouts in burned areas. We
also determined if susceptibility to photoinhibition was influ-
enced by leaf age (gren 1991, Krause et al. 1995).
Materials and methods
Study site and experimental design
The study was carried out in a Quercetum ilicis galloprovin-
ciale (Bols 1962) forest in Puig El Sotarr in the Serra del
Garraf mountains near Barcelona, Spain (4120 N, 155 E;
elevation 663 m). The climate at the site is typically Mediter-
ranean with cold winters, cool wet springs and autumns and
hot dry summers, with a mean annual temperature of 13--
14 C, and an annual precipitation of 500--700 mm. Meteoro-
logical data during the study were obtained from the Vallirana
field station approximately 3 km away from the study site.
The pre-fire vegetation was a 60-year-old forest dominated
by Pinus halepensis Mill. and Quercus ilex. In July 1995, a
wildfire consumed 100 hectares of the forest. In spring 1996,
we set out four plots (100 100 m) in two areas, each area
contained one intact forest (control) plot and one burned plot.
Plant material
Three Quercus ilex root systems in each control plot and four
root systems in each burned plot were randomly selected for
study. The evergreen Q. ilex shrubs were approximately 0.40 m
high in the burned plots and 2.50--3.50 m high in the unburned
plots. In unburned plots, measurements were carried out on old
(CO) and young (CY, newly emerged) leaves, whereas in the
burned plots young leaves of the resprouts (FY) were measured.
Unless otherwise noted, measurements were conducted in
early summer 1996 (late June--July) around midday under
potentially stressful conditions of irradiance, temperature and
vapor pressure deficit (VPD) (Table 1). Unburned and burned
plots from one of the two areas were studied on a given day.
Gas exchange measurements
Net photosynthesis (A) and stomatal conductance (g) were
determined on attached leaves with a portable LI-6200 (Li-
Cor, Inc., Lincoln, NE) system. Each replicate was carried out
in 20--40 s.
Table 1. Climatological data during the study (1996).
Parameter Period 1 Period 2
June 25--July 14 July 15--July 31
Mean daily maximum temp. (C) 28.01 33.35
Mean daily minimum temp. (C) 16.04 20.0
Mean daily temp. (C) 21.39 26.11
Vapor pressure deficit (kPa) 1.25 2.36
Precipitation (mm) 0.18 0.00
TREE PHYSIOLOGY VOLUME 18, 1998
FLECK ET AL.
Chlorophyll fluorescence measurements
Components of chlorophyll fluorescence were quantified in
situ in the forest with a portable modulated fluorometer (Mini-
Pam Photosynthesis Yield Analyzer, Walz, Effeltrich, Ger-
many). The instrument was equipped with a leaf-clip holder
(2030-B; Walz) and a micro-quantum sensor to monitor PAR
and a thermocouple to measure temperature at the lower leaf
surface (Bilger et al. 1995). The fiberoptics axis forms a 60
angle with the leaf plane, avoiding shading of the sample when
external actinic illumination is applied. After clamping the
leaf-clip holder onto the leaf, the actual fluorescence, F, was
monitored to ascertain that it was stable (weak measuring
beam). Maximum fluorescence yield, Fm, was measured dur-
ing a 0.8-s saturating flash at 6000 mol m--2 s--1 and exposure
to natural illumination. The fraction of absorbed light utilized
in electron transport is given by the PSII efficiency or yield
parameter, F/Fm, where Fm is maximal fluorescence in the
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light and F = Fm F (Genty et al. 1989).
After these measurements, one half of the leaf was cut and
immediately frozen and stored in liquid nitrogen for pigment
and soluble proteins determination. The remaining half was
wrapped in aluminum foil to obtain maximum and minimum
dark-adapted fluorescence (Fo, Fm) and Fv /Fm, where Fv =
Fm Fo. The dark-adaptation time was at least 20 min. The
data were corrected for changes in measuring beam irradiance
induced by temperature changes of the Mini Pam. The correc-
tion was calculated by monitoring the fluorescence signal of a
standard provided with the instrument.
Predawn values of Fo, Fm, and Fv /Fm were determined in the
laboratory for the same leaves characterized at midday as
described by Demmig-Adams and Adams (1996). Leaves were
excised underwater and maintained in petri dishes on moist
paper in the laboratory at room temperature and a PAR of about
10 mol m--2 s--1 for 1--3 days until Fv /Fm values were maximal.
Nonphotochemical quenching coefficient (NPQ) described by
(Fm Fm ) /Fm was determined from forest measurements and
from predawn measurements in the laboratory.
Relative water content
Relative water content (RWC) was determined on a sample of
each leaf. Leaf samples were cut underwater and collected in
preweighed tubes containing cooled water and maintained at
4 C. The increase in tube weight was taken as the sample fresh
mass (FM). Tubes were maintained at 4 C for 24 h, and the
leaves weighed after blotting with paper tissue; this gave the
water saturated mass (WSM). Samples were then dried to a
constant weight in a forced-air oven at 60 C to obtain the dry
mass (DM). The RWC was calculated as: RWC = (FM --
DM)100/(WSM -- DM).
Leaf area was determined from photocopies of leaves using
Image Processing System (IPS-200, Kontron, Bildanalyse,
Munich, Germany).
Chemical analyses
Total nitrogen content of the leaves was determined on dried
material ground to a fine powder in a Mixer-Mill 800 (Spex
Industries Inc., Mentuchen, NJ) with balls of tungsten carbide
 PHOTOSYNTHESIS AND PHOTOPROTECTION IN Q. ILEX
and analyzed by gas chromatography (Nitrogen Analyzer
1500, Carlo Erba, Milan, Italy) under standard conditions
(Pella et al. 1984).
Total soluble protein content of the leaves was determined
by the method of Bradford (1976).
Chloroplast pigments were extracted from leaf discs frozen
in liquid N in the field, ground in a mortar with acetone (about
0.3 cm2 of leaf tissue per ml of solvent) in the presence of
sodium ascorbate and analyzed by HPLC as described by
Rivas et al. (1989).
Statistical analyses
Statistical analyses were conducted by multifactor ANOVA
using Statgraphics Plus (Version 2.0, Maringistics Inc.,
Rockville, MD). Significant differences between means were
identified by Bonferronis test, taking P 0.05 as the level of
significance. We tested for interactions between the main ef-
fects (period, treatment and age). For parameters sensitive to
variations in light, PAR was used as a covariate.
The ANOVA revealed that measurements made in the two
areas were not significantly different. There were also no
differences in the measured parameters between the different
root systems of each treatment. As a result, data from both
areas, and from individuals in an area were pooled for sub-
sequent analysis (Winer 1971).
Results
Statistically significant differences were observed between the
dates of measurement; therefore, we distinguished between
two periods. Period 1 included measurements made up to
July 10 and Period 2 included measurements made from
July 10 onward. Environmental parameters during the two
Figure 1. Net photosynthesis (A) in young control leaves (CY), old
control leaves (CO) and resprouts after fire (FY) measured at midday
in Periods 1 and 2. Values are means SE for 10 leaves from 3--4 root
systems per treatment and day. Only leaves of similar orientation in the
trees and exposed to full sun were chosen.
TREE PHYSIOLOGY ON-LINE at http://www.heronpublishing.com
609
periods are shown in Table 1.
Net photosynthesis (A)
During Period 1, A at midday was similar for leaves in all
groups (Figure 1), whereas there was a significant effect of
treatment but not of leaf age during Period 2 (Figure 1). From
Period 1 to Period 2, both young and old control leaves showed
a 75% decline in A, whereas A increased twofold in young
resprouts after fire.
Transpiration (E) and stomatal conductance (g)
Initial values of E and g were similar for leaves in all treat-
ments, whereas there was a significant effect of treatment but
not of age in Period 2 (Figure 2). Both young and old control
leaves showed an 80% decline in g and E in Period 2 compared
with Period 1, whereas young resprouts after fire maintained
similar g and E throughout the study.
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During the measurements made in Period 1, mean VPD, leaf
temperature and PAR in the leaf chamber were 2.4 kPa,
29.5 C and 1323 mol m--2 s--1, respectively, and the corre-
sponding values for Period 2 were 3.4 kPa, 33.7 C and
1435 mol m--2 s--1.
Chlorophyll fluorescence
During Period 1, PSII efficiency (F/Fm) did not differ signifi-
Figure 2. Transpiration (E) and leaf conductance (g) in control young
leaves (CY), control old leaves (CO) and resprouts after fire (FY)
measured at midday in two different periods. Values are mean SE for
ten leaves from 3--4 root systems per treatment and day. Only leaves
of similar orientation in the trees and exposed to full sun were chosen.
610 FLECK ET AL.

and midday Fv /Fm were 0.783 and 0.714, respectively.

For both young and old control leaves, nonphotochemical
quenching (NPQ) increased significantly during the study,
whereas NPQ of young resprouts after fire was relatively
constant (Figure 4). There was a significant effect of leaf age
but not of treatment in Period 1, with young leaves showing
higher values than old leaves. In Period 2, there were both leaf
age and treatment effects on NPQ. Control leaves had higher
NPQ values than resprouts after fire, and young control leaves
had higher NPQ values than old control leaves. Although NPQ
measured in the laboratory (predawn values) showed similar
trends to NPQ measured in the forest, the laboratory estimates
of Fm showed fluctuations, especially during Period 2 (envi-
ronmental conditions were more stressful during Period 2 than
during Period 1); therefore, we used the laboratory data only
to confirm trends observed in NPQ data from the forest.

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Figure 3. Photosystem II efficiency (F/Fm) of young control leaves
(CY), old control leaves (CO) and resprouts after fire (FY) measured
at midday in Periods 1 and 2. For each day of measurement, values are
means SE of seven young and seven old leaves of two control trees
and five leaves of four resprouting trees. Only leaves of similar
orientation in the trees and exposed to full sun were chosen. (The
values on the figure are not corrected for PAR effect; but it was used
as covariate in the statistical treatment.)
cantly between treatments or leaf ages (Figure 3). During
Period 2, there was a significant effect of treatment but not of
leaf age on F/Fm. Highest values of F/Fm were obtained in
young resprouts after fire (0.295), whereas young and old
control leaves had similar values (0.145). Young resprouts after
fire exhibited similar F/Fm values throughout Periods 1 and
2, whereas F/Fm values of both young and old control leaves
declined during the study.
Predawn and midday values of the potential quantum yield
of PSII (Fv /Fm) did not differ significantly between periods,
treatments or leaf ages (Table 2). The mean values of predawn
Zeaxanthin content
During Period 1, zeaxanthin content was dependent on leaf age
but not treatment, with young resprouts after fire and young
control leaves having higher contents than old control leaves
(Figure 5). During Period 2, there were both leaf age and
treatment effects on zeaxanthin content, with control leaves
showing increased values with respect to Period 1. This in-
crease in zeaxanthin content was higher in young control
leaves than in old control leaves. Young resprouts after fire
showed a decline in zeaxanthin content throughout the study.
Zeaxanthin content was closely related to NPQ values, espe-
cially in the young and old control leaves (Figure 6).
Soluble protein and total nitrogen content
Soluble protein content on an area basis showed a significant
leaf age effect but not a treatment effect (Figure 7). The protein

Table 2. Predawn and midday potential quantum yield of PSII

(Fv /Fm) of Quercus ilex leaves in Periods 1 and 2. For each day of
measurement, values are means of seven young and seven old leaves
of two control trees and fives leaves of four resprouting trees. All SE
values were less than 0.01. Abbreviations: CY = young control leaves;
CO = old control leaves; and FY = young resprouts after fire.

Period 1 Period 2
June 28 July 3 July 9 July 23 July 26 July 30
Predawn Fv /Fm
CY 0.789 0.797 --1 -- 0.786 0.79
CO 0.759 0.783 -- -- 0.792 0.766
FY 0.744 0.79 -- -- 0.803 0.734
Midday Fv /Fm
CY 0.712 0.727 0.725 0.727 0.699 0.695 Figure 4. Nonphotochemical quenching (NPQ) in young control
CO 0.704 0.754 0.687 0.673 0.705 0.707 leaves (CY), old control leaves (CO) and resprouts after fire (FY)
FY 0.732 0.716 0.701 0.765 0.692 0.723 measured at midday in Periods 1 and 2. For each day of measurement,
values are means SE of seven young and of seven old leaves of two
1
No data available. control trees and five leaves of four resprouting Holm oaks.

TREE PHYSIOLOGY VOLUME 18, 1998

 PHOTOSYNTHESIS AND PHOTOPROTECTION IN Q. ILEX
Figure 5. Zeaxanthin content (Z x 100)/(V + A + Z) in young control
leaves (CY), old control leaves (CO) and resprouts after fire (FY)
measured at midday in Periods 1 and 2. Abbreviations: V = violaxan-
thin, A = antheraxanthin. Values are means SE of 10--20 individual
leaves of 2--4 different root systems per treatment, age and day.
contents of young resprouts after fire and young control leaves
were similar and did not vary from Period 1 to Period 2,
whereas old control leaves had higher protein contents in
Period 1 and lower protein contents in Period 2 than young
leaves. Nitrogen content on an area basis also showed a leaf
age effect but not a treatment effect, increasing significantly
from Period 1 to Period 2 in young leaves in both treatments
(Figure 7). Nitrogen content in old control leaves was similar
in both periods.
Relative water content and leaf dry mass per area (LMA)
In all leaves measured, RWC decreased slightly but signifi-
cantly from 89% in Period 1 to 81% in Period 2. Within each
period, there were no significant effects of treatment or leaf age
on RWC. There was a leaf age effect on LMA but not a
treatment effect. The LMA in old control leaves was similar in
Periods 1 and 2, whereas in young resprouts after fire and
young control leaves, LMA increased significantly from Pe-
riod 1 to Period 2.
Discussion
Several physiological characteristics of the resprouting vege-
tation of Quercus ilex after fire were significantly different
from the original vegetation, especially under the stressful
conditions that occur in a Mediterranean climate in the sum-
mer. The fire treatment effect was enhanced when the plants
were subjected to increased temperature, irradiance and VPD
at midday.
At the beginning of the study (Period 1), gas exchange and
PSII efficiency at midday did not differ significantly between
old and young leaves from control (unburned) plants and
resprouts that appeared after fire (Figures 1--3). In Period 2,
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611
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Figure 6. Relationship between nonphotochemical quenching (NPQ)
and zeaxanthin content (Z x 100/(V + A + Z) in old control leaves
(CO), young control leaves (CY) and resprouts after fire (FY) meas-
ured at midday. Abbreviations: V = violaxanthin, A = antheraxanthin.
when conditions were stressful (see Table 1), fire treatment had
a significant effect on gas exchange and PSII efficiency,
whereas there was no effect of leaf age on these parameters.
Young resprouts after fire did not show midday depressions
of photosynthesis and leaf conductance (Figures 1 and 2),
whereas both young and old control leaves showed marked
midday declines in both A and g. Increases in A and g in
burned plots with respect to unburned plots in hot seasons
have been reported previously (e.g., Knapp 1985, Hastings et
al. 1989, Reich et al. 1990, Busch and Smith 1993, Fleck et
al. 1995, 1996b). The absence of differences in A and g
between old and young leaves of control plots indicates that
leaf ontogeny does not play a determinant role in the differ-
ences observed between pre- and post-burn vegetation ( Fleck
et al. 1995).
In Period 2, the relative water content (RWC) decreased
slightly but significantly in all leaves with respect to Period 1
(Figure 8). Quercus ilex is an evergreen sclerophyllous deep-
rooted species and RWC can be quite stable throughout the
year (Rhizopoulou and Mitrakos 1990). Despite a nearly four-
fold higher transpiration rate in young resprouts after fire in
Period 2 than in Period 1 (Figure 2), RWC of resprouts re-
mained higher than that of control leaves. Increased water
612 FLECK ET AL.
Figure 7. Total soluble protein and nitrogen contents in young control
leaves (CY), old control leaves (CO) and resprouts after fire (FY) in
Periods 1 and 2. Values are means SE of 3--4 replicates from 5--10
leaves from 2--4 different root systems per treatment, age and day.
availability from the extensive root system to the small re-
sprouting shoots (De Souza et al. 1986, Saruwatari and Davis
1989) or increased soil-to-leaf hydraulic conductivity (Kruger
and Reich 1993) could account for this finding. Moreover,
cooling is induced by this non-conservative water use (Busch
and Smith 1993). Increased stomatal opening in the resprouts
under conditions that favor water stress contributed to the
observed increases in CO2 fixation and growth during Period 2.
In Period 2, nitrogen and soluble protein contents were
affected by leaf age but not treatment. Although the nitrogen
and soluble protein contents of young leaves of burned and
unburned plots were similar (Figure 7), photosynthetic rates
Figure 8. Relative water content (RWC) and leaf mass per area (LMA)
in young control leaves (CY), old control leaves (CO) and resprouts
after fire (FY) in Periods 1 and 2. Values are means SE of 10--20
leaves per treatment, age and day.
TREE PHYSIOLOGY VOLUME 18, 1998
were much lower in young leaves from the unburned site
indicating stomatal limitation. During Period 2, young leaves
from unburned and burned plots increased their N content but
maintained their soluble protein content with respect to Pe-
riod 1. Data on chloroplast pigments on an area basis (not
shown) indicated that the increased N in Period 2 was invested
in thylakoids of the young leaves from unburned and burned
plots.
In vegetation from the unburned plot, the decrease in midday
photochemical efficiency of PSII (F/Fm) in Period 2 (Fig-
ure 3) reflects a decrease in the fraction of absorbed light
energy utilized in photochemical reactions and the participa-
tion of protection mechanisms against excess solar energy
(Genty et al. 1989). During midday depression of gas ex-
change, the effects of high temperatures and the reduction in
the internal CO2 concentration as a result of stomatal closure
may favor the oxygenase activity of Rubisco thereby leading
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to increased photorespiration, which may also play a role in
protection against photodamage (Brestic et al. 1995, Kozaki
and Takeba 1996, Lovelock and Winter 1996). Our results on
PSII efficiency (Figure 3) indicate that photorespiration or
cyclic electron transport may not have been sufficient to main-
tain high electron transport rates in vegetation from the un-
burned plot as reported by Epron et al. (1993) for oak species.
We did not test the protective role of antioxidant systems in
leaves, but they have been reported to increase in Quercus
suber L. during the Mediterranean summer (Faria et al. 1996).
The increase in NPQ values to around three in both young
and old control leaves during Period 2 (Figure 4) indicates
increased thermal energy dissipation in the antennae. Down-
regulation of photosynthetic electron transport associated with
an increase in nonphotochemical quenching of chlorophyll
fluorescence has been described by Sharkey et al. (1986) and
Weis and Berry (1987). A process involved in NPQ is the
conversion of violaxanthin to antheraxanthin and zeaxanthin,
which mediates the harmless dissipation of excess energy as
heat (Demmig and Bjrkman 1987). As reported for other
species (Demmig-Adams and Adams 1996), we observed a
good correlation between zeaxanthin content and NPQ in both
control leaves and resprouts (Figure 6).
Under stressful conditions at midday in Period 2, PSII effi-
ciency remained substantially higher and NPQ lower in young
resprouts after fire compared with control values. In the re-
sprouts after fire, higher stomatal conductance and N content,
which favored carbon fixation, created a higher photochemi-
cal sink (as defined by Osmond et al. 1997) for photons.
However, in the control vegetation, CO2 fixation was severely
restricted by reduced stomatal conductance, and the thermal
sink created by nonphotochemical processes was enhanced.
Even at midday, resprouts after fire did not show enhanced
non-radiative dissipation as measured by NPQ. This finding
supports the conclusion of Demmig-Adams et al. (1992) that
NPQ (thermal dissipation in antennae) accounts for the light
energy in excess of that used in photochemistry. The greater
photochemistry of the resprouts would make a higher thermal
sink unnecessary.
Observed values of predawn potential quantum yield of PSII
(Fv /Fm) (Table 2) were within the range (0.75--0.85) that is
 PHOTOSYNTHESIS AND PHOTOPROTECTION IN Q. ILEX
normal for well-watered plants (Bjrkman and Demmig
1987). Midday Fv /Fm values were only slightly lower than
predawn Fv /Fm values, indicating that none of the leaves were
photoinhibited in either period. Mthy et al. (1996) reported
that Fv /Fm of Quercus ilex was only affected when leaf pre-
dawn water potential was less than --4 MPa, a value rarely
observed in mature trees in the field.
The decrease in photosynthetic rate at midday in control
leaves was not paralleled by a decrease in Fv /Fm indicating that
the xanthophyll cycle is an effective way of dissipating energy
and thereby avoiding PSII damage in these Quercus ilex
leaves. We suggest that low stomatal conductance is the pri-
mary limiting factor in Quercus ilex leaves, initiating regula-
tion of carboxylation activity and energy dissipation
processes.
Acknowledgments
This work was supported by funds from DGYCIT (PB94-0930).
Participation by K.P. Hogan was supported by Comissionat per a
Universitat i Recerca, Generalitat de Catalunya (Estada de professors-
investigadors visitants 1995). We thank Direcci General de Qualitat
Ambiental, Departament de Medi Ambient, Generalitat de Catalunya
for the climatological data, J. Caas and J. Vilam of Parc Natural de
Collserola for place facilities and Servei Cientfic-Tcnic de la UB for
technical assistance.
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