Tree Physiology

14,725-133
0 1994 Heron Publishing-Victoria,

Canada

Relationship between COz-dependent 02 evolution and

photosystem II activity in oak (@emus petraea) trees grown in
the field and in seedlingsgrown in ambient or elevated CO2
DANIEL EPRON,1,2,3 ERWIN DREYER,
MARC GUEHL]
Laboratoire

d Ecophysiologie

2 Present

address:

3 Author

to whom

Received

September

Laboratoire
correspondence

Forest&e,
Ecologic
should

Centre
V&&ale,

CATHERINE
INRA

de Nancy,

Universite

Paris

PICON and JEAN

54280

Champenoux,

Sud, 91405

Orsay

France
cedex,

France

be sent

24, 1993

Summary
The light-response
of the apparent quantum yield of photosynthetic
02 evolution
(QoJ under non-photorespiratory
conditions
was measured
together with the photochemical
efficiency
of PS II @F/F,,,),
the
photochemical
efficiency
of open PS II reaction
centers (F/F,,,)
and the photochemical
fluorescence
quenching
(qr) of leaf disks punched from oak leaves of seedlings grown in ambient (350 ymol mol-)
or elevated (700 pmol mol-)
CO2 in a greenhouse,
and from sunlit leaves of mature oak trees (Quercus
petraea
(Man)
Liebl.). There were marked differences
between seedlings
and trees. In seedlings,
CO2
concentration
during growth did not modify
the light response of photosynthesis
or PS II activity.
There
was a single linear relationship
between Qo2 and AFIF,
in seedling leaves that was independent
of the
CO2 concentration
imposed during growth. In contrast, this relationship
was curvilinear
in sunlit leaves
of adult trees. In seedling leaves, the decrease in qp (i.e., the proportion
of open PS II reaction centers)
largely accounted
for the decrease in AF/F ,,,, whereas the decrease in AFIF,,, in sunlit leaves of mature
oak trees was dependent on both qp and F,/F,.
Keywords:

chlorophyll

fluorescence,

gas exchange,

photochemical

efficiency,

photosynthesis.

Introduction
Simultaneous determinations of in viva fluorescence and leaf gas exchange (CO2
assimilation or 02 evolution) have been used to calculate the rate of light-driven
photosynthetic electron flow and to estimate its partitioning among assimilative and
dissipative processes (Peterson 1989, Comic and Briantais 1991).
Two models have been used to describe the relationship between chlorophyll
fluorescence and the quantum yield of electron transport. Weis and Berry (1987)
found that the quantum yield of electron transport was linearly related to the
non-photochemical quenching of fluorescence when divided by the photochemical
quenching of fluorescence, and Genty et al. (1989) found that it was also related to
the product of the photochemical quenching and the quantum yield of open reaction
centers as determined by F,/F,, the ratio of variable to maximal fluorescence (Butler
1978). This second model provides a useful tool for ecophysiological studies because
it does not require the measurement of initial fluorescence (Fo), which is difficult to
obtain, particularly in natural conditions.

126

EPRON,

DREYER,

PICON

AND

GUEHL

However, the universality of both models has been questioned because non-linearity is often observed (Peterson et al. 1988, Seaton and Walker 1990, Oquist and Chow
1992). It has been suggested that the observed discrepancies may reflect differences
among species or growth conditions (Peterson et al. 1988). Recently, Evans et al.
(1993) have shown that curvilinearity is observed if the gradients in light absorption
and photosynthetic capacity are not identical.
Relationships between the quantum yield of photosynthetic oxygen evolution and
the photochemical efficiency of PS II were established for sunlit leaves of adult trees
and for leaves of seedlings grown in ambient (350 pmol mol-) or elevated (700
pmol mol-) CO2 in a greenhouse. We studied possible effects of growth conditions
on these relationships and on the contribution of non-photochemical energy dissipation to the modulation of PS II activity.

Material

and methods

Plant material and experimental conditions

Seedlings of Quercus petraea (Matt.) Liebl. (seed origin, Saint Dizier forest, eastern
France) were grown in two polyethylene chambers (3 x 4 m2 area, 2.3 m height) in
a naturally illuminated greenhouse. One of these chambers was maintained at
ambient CO2 molar fraction (350 pmol mol-), and the other one at twice the ambient
CO;! molar fraction (700 pmol mol-). Seedlings were grown individually in 5-liter
pots filled with a l/l (v/v) mixture of sand and blond sphagnum peat fertilized with
20 g of a slow release fertilizer (Nutricote 100, N,P,K, 13,13,13) supplemented with
a mixture of oligoelements. Plants were kept well watered by irrigating them twice
a week. Photosynthetic photon flux density inside the chambers was one third of that
recorded outside, with maximal values around 500 pmol mm2 s-. A complete
description of the experimental design is given in Guehl et al. (1994).
In addition, thirty-year-old Quercus petraea trees growing in a stand in Champenoux forest near Nancy, France were also studied. They were kept well watered
throughout the summer (see Epron et al. 1992 for details).
In August 1992, measurements were made on well-expanded leaves taken either
from the sun-exposed part of the crown of two dominant trees (three leaves per tree)
or from the last fully developed growth flush of six seedlings (4 months old) per CO2
treatment. Seedlings were darkened at least 12 h before excision of the leaf, and
leaves were taken before dawn on trees grown outdoors. A leaf disk (10 cm2) was
immediately punched from each of the excised leaves and enclosed in the electrode
chamber.

Photosynthetic oxygen evolution and chlorophyllfluorescence

measurements

Measurements of photosynthetic oxygen evolution rates and chlorophyll fluorescence yield were made at 25 C in a leaf disk electrode chamber (LD2, Hansatech,
Kings Lynn, U.K.) filled with water-saturated air containing 5% CO2 and 19% 02.
The leaf was illuminated from the top of the chamber by a halogen lamp (LS2,

FLUORESCENCE

AND

PHOTOSYNTHESIS

IN OAK

121

Hansatech, Kings Lynn, U.K.). Neutral density filters were used in combination to
obtain incident photon flux densities (PFD) ranging from 47 to 1142 pmol me2 s-l.
Chlorophyll fluorescence was measured with a modulated fluorometer (PAM 101,
Walz, Effeltrich, Germany). Both the measuring modulated red light (0.1 pmol rnp2
S -, 100 kH) and the flashes of saturating white light (6000 pmol rnp2 s-l, 800 ms
duration), delivered by a halogen source (KL1500, Schott, Germany), were provided
to the leaf disk by a multi-branched fiber optic at an angle of 45 to the top of the
chamber. Fluorescence was recorded by another branch of the fiber optic.
Before fluorescence measurements, the leaf disks were darkened for 15 min in the
presence of 5% CO2. Initial fluorescence (Fa), when all reaction centers are open,
was recorded using the measuring modulated red light. Maximal fluorescence (F,),
when all reaction centers are closed and all non-photochemical fluorescence quenchings are relaxed, was recorded during imposition of a flash of saturating white light.
The leaf disk was then illuminated until a steady state rate of 02 evolution was
achieved (30 to 50 min). Steady state fluorescence yield (F) and maximal fluorescence yield following a flash of saturating white light (F,) were recorded. Minimal
fluorescence (Fo) and dark respiration were recorded during dark periods (5 min)
between two successive PFD values. Because no dark respiration was detected, the
rate of 02 evolution was used directly as the rate of photosynthetic 02 evolution.
Photochemical fluorescence quenching (qJ was calculated according to Bilger and
Schreiber (1986) as qp = (F, - F)/(F,,, - Fa) and used as an estimate of the fraction
of open reaction centers. The photochemical efficiency of PS II (m/F,)
and of open
PS II reaction centers (F,/F,) were estimated as F,/F, = 1 - (Fo/Fm) (Butler
1978) and AF/F, = 1 - (FIF,) = qp F,/F, (Genty et al. 1989).
The apparent quantum yield of 02 evolution (Q$ was calculated as the ratio of
02 evolution rate to the corresponding incident PFD. After completion of a light
response curve by increasing PFD step by step, chlorophyll content was determined.
A small part of the leaf disk (3.15 cm2) was extracted in 10 ml of dimethylsulfoxide
at 65 C for 90 min (Hiscox and Israeltam 1979). The remaining part of the leaf disk
was weighed after oven-drying at 60 C for 48 h, and dry weight was used to
calculate specific leaf area (SLA).
Results
The responses of the rate of photosynthetic oxygen evolution to photon flux density
(PFD) are shown in Figure 1 (upper panel). Although the difference was not
statistically significant, slightly higher rates of 02 evolution were observed in
seedlings grown in elevated CO2 than in seedlings grown in ambient CO2 at PFD
values above 600 pmol m-* s-. At lower u-radiances, no differences in rates of 02
evolution were observed between seedlings grown in ambient or elevated CO2
(one-way ANOVA, P = 0.05). At all PFD values, sunlit leaves from mature trees
exhibited lower rates of 02 evolution than leaves from seedlings.
Despite a significantly lower specific leaf area (SLA, Table l), leaves of seedlings
grown in elevated CO2 had a similar chlorophyll content to seedlings grown in

728

EPRON,

DREYER,

PICON

AND

GUEHL

Figure 1. Responses
of photosynthetic
02 evolution
(pm01 m -2 s-) and photochemical
efficiency
of
PS II (A/W,,,)
to incident photon flux density (PFD, pmol m -2 s-) in leaves of oak seedlings
grown in
ambient (0) or elevated (0) CO2 and in sunlit leaves of adult oak trees (A). Mean values and standard
deviations
are given for six replicates.
Measurements
were carried out in 5% CO2 at 25 C.

ambient CO2. As a consequence, maximal rates of photosynthetic 02 evolution

expressed on a chlorophyll content basis were very similar (Table 1). Sunlit leaves
of trees grown in natural conditions exhibited lower SLA and higher chlorophyll
content than seedlings grown in ambient COZ, but a significantly lower maximal rate
of photosynthetic 02 evolution when expressed on a leaf area basis or on a chlorophyll content basis (Table 1).
Increases in PFD led to a similar decline in the photochemical efficiency of PS II
(M/F,,,) for leaves of seedlings grown in ambient or elevated CO2 (Figure 1, lower
panel). At any given PFD, and despite lower rates of 02 evolution, sunlit leaves of
Table 1. Specific
leaf areas (SLA, dm gg), chlorophyll
contents
(mg dme2) and maximal
rates of
on a leaf area basis (pmol m-2 ss) or on a chlorophyll
photosynthetic
02 evolution
(02 ,,,) expressed
content basis (pm01 g&
s-) in leaves of oak seedlings
grown at ambient (350) or elevated (700 pmol
mol-)
CO2 and in sunlit leaves of adult oak trees. Mean values and standard deviations
are given for six
replicates.
Values followed
by the same letter are not significantly
different
(one-way
ANOVA,
P = 0.05).
Treatment

Seedlings grown at
ambient CO2
Seedlings
grown at
elevated CO2
Trees grown outdoors

SLA
(dm2 g-)

[Chlorophyll]
(mg dmm2)

02max

2.04 f 0.12a

5.95 k 0.28a

20.2 f 3.5ab

34.0 * 5.5a

1.75 f 0.15b

6.34 k0.37a

23.8 + 5.9a

35.5 k 9.7a

0.94 k 0.08~

8.36 k 0.36b

16.7 f 3.5b

20.1 f 4.4b

(pm01

02max

m -2 s- 1

@mOl

gchl -s-3

FLUORESCENCE

AND

PHOTOSYNTHESIS

IN OAK

729

trees exhibited slightly higher AF/F, values than seedling leaves.

Figure 2 shows the linear relationship between AF/F, and the apparent quantum
yield of photosynthetic 02 evolution (QoJ of seedling leaves. The linear regression
was similar for both treatments, whereas the relationship was curvilinear for sunlit
leaves of adult trees. Increasing PFD led first to a large decrease in iDo, without any
significant change in AF/Fm, followed by a steep decrease in AFIF,,, at low Qo,, i.e.,
at PFD values higher than 250 pmol mm2 s-l.
In leaves of oak seedlings, there was a close linear relationship between photochemical fluorescence quenching (qr) and AF/Fm (Figure 3, upper panel). The
photochemical efficiency of open PS II reaction centers (F,/F,) remained largely
unaffected when AF/F, decreased from 0.7 to 0.3, but decreased at lower values of
AFIF, (Figure 3, lower panel). In sunlit leaves, the decrease in AFIF, was linearly
related to decreases in both qp and F,/F,.

Discussion
Photosynthetic responses of oak seedlings to elevated CO2
Long-term exposure to elevated CO2 often leads to a reduction in the photosynthetic
capacity of leaves, especially in tree species (DeLucia et al. 1985, Kaushal et al.
1989, Mousseau and Saugier 1991, Bunce 1992), but a lack of down regulation of
photosynthetic capacity or even a positive acclimation has also been observed
(Bunce 1992, Curtis and Teery 1992, Pettersson and McDonald 1992). In our
experiment, oak seedlings showed no down regulation of photosynthetic capacity in
response to long-term exposure to elevated CO*. Both chlorophyll content per unit
area and maximal rate of photosynthetic oxygen evolution per chlorophyll content
were similar in seedlings grown at ambient and at elevated CO2. The similarity in the

0.6

04

0.2

n.n
-.0.00
t&u&urn

0.02

0.04

0.06

yield Imolt

0.08
mol.I

0.10
photon)

Figure
2. Relationships
between
the quantum
yield of photosynthetic
02 evolution
(mol 02 mol
photon-)
and photochemical
efficiency
of PS II (W/F,,,)
for leaves of oak seedlings grown in ambient
(0) or elevated (0) CO2 and in sunlit leaves of adult oak trees (A). Measurements
were carried out in
5% CO2 at 25 C at various incident photon flux densities ranging from 47 to 1142 pmol mm2 s-. Solid
lines are linear regressions
fitted through
the data (y = -0.045
+ 7.574x, r2= 0.948 and y = -0.043
+
8.004x, 2 = 0.935 for seedlings
grown in ambient and elevated CO2, respectively).

EPRON,

0,o

0.2

0,4

0,6

DREYER,

PICON

AND

GUEHL

0,s

AFIFIII

Figure 3. Relationships
between
the photochemical
efficiency
of PS II (W/Fm)
and either the photochemical
fluorescence
quenching
(qr) or the photochemical
efficiency
of open PS II reaction
centers
(F,/F,)
for leaves of oak seedlings
grown in ambient (0) or elevated (0) CO2 and in sunlit leaves of
adult oak trees (A). Measurements
were carried out in 5% CO2 at 25 C at various incident photon flux
densities ranging from 47 to 1142 ymol m -2 s-. Solid lines are linear regressions
fitted through the data:
for seedlings,
the decrease
in AFIF ;n was linearly
related to a decrease in qp (J = 0.06 1 + 1.425x, r2 =
0.987 andy = 0.082 + 1.422x, r2 = 0.992 for seedlings grown in ambient and elevated CO2, respectively),
and for sunlit leaves, the decrease in AF/F,
was linearly related to a decrease in both qp (y = 0.332 +
1.005x, r2 = 0.942) and F/F,,, CJ= 0.232 +0.690x,
r2 = 0.955).

light response curves of photosynthetic oxygen evolution rates and PS II photochemical efficiency of seedlings in the two CO2 treatments suggests that, under non-photorespiratory conditions, elevated CO2 has no effect on photosynthetic capacity and
quantum yield of CO2 fixation. Guehl et al. (1994) reported that, under the same
growth conditions, the biomass accumulation of Quercus petruea seedlings in
elevated CO;? was 2.4 times higher than that observed in ambient CO*. The absence
of down regulation of photosynthetic capacity in response to elevated CO2 is
consistent with the marked COZ-induced stimulation of growth.

Relationship between quantum yield of photosynthetic oxygen evolution and PS II

photochemistry
Under non-photorespiratory conditions, there was a linear relationship between the
quantum yield of photosynthetic oxygen evolution and the product of the efficiency
of excitation capture by open PS II reaction centers (F,/F,,,) and the fraction of open
centers (qJ, indicating that AFIF,,, represents the quantum yield of charge separation
at PS II, which is linearly correlated with the quantum yield of gross photosynthetic
O2 evolution under non-photorespiratory
conditions, i.e., the quantum yield of the
light-induced linear electron flow (Genty et al. 1989). Similar results have been

FLUORESCENCE AND PHOTOSYNTHESIS IN OAK

731

obtained in other Cs species (Keiller and Walker 1990, Krall et al. 1991, Plesnicar
and Pankovic 199 1, Comic and Briantais 199 1). The relationship between apparent
quantum yield of photosynthetic oxygen evolution and PS II photochemical efficiency was similar in seedlings grown in ambient and elevated CO*, indicating
similar leaf absorbance and energy partitioning between PS II and PS I in the two
CO:! treatments, because this slope is mainly determined by these two parameters
(Genty et al. 1989). The slope values (around 8) are similar to those reported by
Genty et al. (1989) in barley leaves and by Comic and Briantais (1991) in bean
leaves.
Sunlit leaves of adult trees only exhibited a linear relationship between quantum
yield of photosynthetic 02 evolution and the photochemical efficiency of PS II at low
quantum yields of photosynthetic oxygen evolution in the range of 0.01 to 0.035.
Similar observations have been reported by Seaton and Walker (1990) and Oquist
and Chow (1992) for other Cs species. Recently, Evans et al. (1993) showed that
curvilinearity is observed if gradients in light absorption and photosynthetic capacity
are not identical. Because sunlit leaves of trees had lower specific leaf area and so
probably higher leaf thickness than leaves of seedlings, this explanation may also
account for the observed curvilinearity in our study.
Modulation of PS II photochemical efficiency
The quantum yield of PS II electron transport is determined by both the efficiency of
open PS II reaction centers and the proportion of open PS II reaction centers (Genty
et al. 1989). The relative contribution of each process to the decrease in AF/F, in
response to an increase in irradiance differed between leaves of seedlings and trees.
In seedling leaves, the decrease in qp largely accounted for the decrease in AF/F,,
whereas in tree leaves both processes contributed to the reduction in AFIF,. A
decrease in F,IF,, i.e., an increase in non-photochemical deactivation of PS II, is
often related to an increase in zeaxanthin content (Demmig Adams et al. 1989, Bilger
and Bjorkman 1990, Gilmore and Yamamoto 1991). Our results confirm the findings
of Demmig et al. (1987) and Thayer and Bjorkman (1990) of a higher capacity for
light-induced zeaxanthin accumulation in sun than in shade leaves.
Non-photochemical deactivation of PS II had a greater influence in the modulation
of PS II electron transport in sunlit leaves of trees than in leaves of seedlings. This
may represent a physiological adaptation to higher irradiances during leaf ontogeny.
In contrast, elevated CO2 did not modify the light responses of photosynthesis and
PS II activity of oak seedlings grown under limiting light in a greenhouse.
Acknowledgments
Audrey Lallemant and FrCdCrique Lafon are acknowledged for their skillful technical assistance.
References
Bilger, W. and 0. BjGrkman. 1990. Temperature dependence of violaxanthin de-epoxidation and nonphotochemical fluorescence quenching in intact leaves of Gossypium
hirsutum
L. and Malva
parviJlora
L. Planta 184:226-234.

732

EPRON,

DREYER,

PICON

AND

GUEHL

Bilger, W. and U. Schreiber.

1986. Energy-dependent
quenching
of dark level chlorophyll
fluorescence
in intact leaves. Photosynth.
Rea. 10:303-308.
Bunce, J.A. 1992. Stomatal
conductance,
photosynthesis
and respiration
of temperate
deciduous
tree
seedlings
grown
outdoors
at an elevated
concentration
of carbon
dioxide.
Plant Cell Environ.
15541-549.
Butler, W.L. 1978. Energy distribution
in the photochemical
apparatus
of photosynthesis.
Annu. Rev.
Plant Physiol.
29:345-378.
Comic, G. and J.M. Briantais.
1991. Partitioning
of photosynthetic
electron
flow between CO2 and 02
reduction
in a Cs leaf (Phase&s
vulgaris
L.) at different
CO2 concentrations
and during drought
stress. Planta 183:255-260.
Curtis, P.S. and J.A. Teery. 1992. Seasonal responses
of leaf gas exchange
to elevated carbon dioxide
in
Populus grandidentata.
Can. J. For. Res. 22: 1320-1325.
DeLucia,
E.H., T.W. Sasek and B.R. Strain. 1985. Photosynthetic
inhibition
after long-term
exposure
to
elevated levels of atmospheric
carbon dioxide. Photosynth.
Res. 7: 175-184.
Demmig
Adams, B., K. Winter, A. Krtiger and EC. Czygan.
1989. Light response of CO2 assimilation,
dissipation
of excess excitation
energy, and zeaxanthin
content of sun and shade leaves. Plant Physiol.
90:881-886.
Demmig,
B., K. Winter, A. Kruger and EC. Czygan.
1987. Photoinhibition
and zeaxanthin
formation
in
intact leaves. A possible role of the xanthophyll
cycle in the dissipation
of excess light energy. Plant
Physiol. 84:218-224.
Epron, D., E. Dreyer and N. Bmda. 1992. Photosynthesis
of oak trees [Quercus
pefruea
(Matt.)
Liebl.]
during drought
under field conditions:
diurnal course of net CO2 assimilation
and photochemical
efficiency
of photosystem
II. Plant Cell Environ.
15:809-820.
Evans, J.R., I. Jakobson
and E. Ggren.
1993. Photosynthetic
light response.
2. Gradients
of light
absorption
and photosynthetic
capacity.
Planta 189: 191-200.
Genty, B., J.M. Briantais
and N.R. Baker.
1989. The relationship
between
the quantum
yield of
photosynthetic
electron transport
and quenching
of chlorophyll
fluorescence.
Biochim.
Biophys.
Acta
990:87-92.
Gilmore,
A.M., and H.Y. Yamamoto.
1991. Zeaxanthin
formation
and energy-dependent
fluorescence
quenching
in pea chloroplasts
under artificially
mediated
linear and cyclic electron
transport.
Plant
Physiol. 96:635-643.
Guehl, J.M., C. Picon, G. Aussenac
and P Gross. 1994. The interactive
effects of elevated CO2 and soil
drought
on growth
and transpiration
efficiency
and its determinants
in two European
forest tree
species. Tree Physiol.
14:707-724.
Hiscox, J.D. and G.F. Israeltam.
1979. Amethod
for the extraction
of chlorophyll
from leaftissue
without
maceration.
Can. J. Bot. 57:768-776.
Kaushal,
P, J.M. Guehl and G. Aussenac.
1989. Differential
growth
response
to atmospheric
carbon
dioxide enrichment
in seedlings of Cedrus atlantica
and Pinus nigru ssp. Laricio var. Corsicana.
Can.
J. For. Res. 19:13.51-1358.
Keiller, D.R. and D.A. Walker. 1990. The use of chlorophyll
fluorescence
to predict CO2 fixation during
photosynthetic
oscillations.
Proc. Roy. Sot. Lond. B 241:59-64.
Krall, J.P., G.E. Edwards
and M.S.B. Ku. 1991. Quantum
yield of photosystem
II and efficiency
of CO2
fixation
in Flaveriu
(Asteraceae)
species under varying
light and COz. Amt.
J. Plant Physiol.
18:369-383.
Mousseau,
M. and B. Saugier. 199 I. The direct effect of increased
CO2 on gas exchange
and growth of
forest tree species. J. Exp. Bot. 43:1121-1130.
Oquist,
G. and W.S. Chow. 1992. On the relationship
between
the quantum
yield of photosystem
II
electron
transport,
as determined
by chlorophyll
fluorescence
and the quantum yield of COz-depcndent 02 evolution.
Photosynth.
Res. 33:5 l-62.
Peterson,
R.B. 1989. Partitioning
of noncyclic
photosynthetic
electron
transport
to 02 dependent
dissipative
processes
as probed by fluorescence
and CO2 exchange.
Plant Physiol. 90:1322-1328.
Peterson,
R.B., M.N. Sivak and D.A. Walker. 1988. Relationship
between steady state fluorescence
yield
and photosynthetic
efficiency
in spinach leaf tissue. Plant Physiol.
88: 158-163.

FLUORESCENCE

AND

PHOTOSYNTHESIS

IN OAK

133

Pettersson,
R. and A.J.S. McDonald.
1992. Effects of elevated carbon dioxide concentration
on photosynthesis
and growth
of small birch plants (Betula pendulu
Roth.) at optimal
nutrition.
Plant Cell
Environ.
15:911-919.
Plesnicar,
M. and D. Pankovic.
1991. Relationship
between chlorophyll
fluorescence
and photosynthetic
02 evolution
in several Helianthus
species. Plant Physiol. B&hem.
29:681-688.
Seaton, G.G.R. and D.A. Walker. 1990. Chlorophyll
fluorescence
as a measure of photosynthetic
carbon
assimilation.
Proc. Roy. Sot. Lond. B 242:29-35.
Thayer,
S.S. and 0. Bjorkman.
1990. Leaf xanthophyll
content
and composition
in sun and shade
determined
by HPLC. Photosynth.
Res. 23.:331-343.
Weis, E. and J.A. Berry. 1987. Quantum
efficiency
of photosystem
II in relation to energy-dependent
quenching
of chlorophyll
fluorescence.
Biochim.
Biophys.
Acta 894:198-208.