Tree Physiology 21, 1039–1045
© 2001 Heron Publishing—Victoria, Canada
Gas exchange characteristics of a Canarian laurel forest tree species
(Laurus azorica) in relation to environmental conditions and leaf
canopy position
AGUEDA M. GONZÁLEZ-RODRÍGUEZ, DOMINGO MORALES and M. SOLEDAD JIMÉNEZ
Dpto. Biología Vegetal, Facultad de Farmacia, Universidad de La Laguna, E-38207 La Laguna, Tenerife, Spain
Received October 16, 2000
Summary Diurnal courses of gas exchange were measured
over a 1-year period in fully expanded current-year leaves in
the upper (sun-exposed, 18 m above ground) and the lower
(shaded, 12 m above ground) canopy of Laurus azorica (Seub.)
Franco, a major canopy species of the Canarian laurel forest in
Tenerife, Canary Islands, Spain. Laurus azorica exhibited high
leaf plasticity in gas exchange characteristics, with a maximum
carbon assimilation rate (Amax) of shade leaves about 50% that
of sun leaves. This difference reflects the high leaf area index
(LAI) of the stand and the correspondingly sharp light attenua-
tion with increasing canopy depth. In sun leaves, Amax peaked
at about 11 µmol m –2 s –1 and maximum transpiration (E) was
about 8 mmol m –2 s –1, which corresponded with a maximum
stomatal conductance (gs) of about 650 mmol m –2 s –1. Mean
maximum instantaneous water-use efficiency (WUE) was
1.5 mmol mol –1 and the mean maximum A/gs was 20–35 µmol
mol –1. Mean minimum internal CO2 concentration (Ci ) was
225 µmol mol –1. Although high air vapor pressure deficit
(VPD) caused a small decrease in gs, it remained high enough
to maintain relatively high A and E. These gas exchange char-
acteristics indicate a non-conservative use of water, which is
appropriate for a species subject to droughts that are mild or of
short duration. In this respect, Laurus azorica differs from its
congener, L. nobilis L., of the Mediterranean region and other
shrubs growing in Mediterranean-type climates in California
and Chile that have to withstand more severe or more pro-
longed droughts.
Keywords: carbon assimilation rate, evergreen broad-leaved
forest, shade leaves, stomatal conductance, sun leaves, wa-
ter-use efficiency.
Introduction
The Canarian laurel forest is thought to be a relic of the Ter-
tiary Mediterranean flora that occupied southern Europe and
northern Africa about 20 million years ago. Climatic changes
during the Quaternary period resulted in conditions suitable
for the survival of this forest type at scattered locations around
the world. Among these is the most humid portion of the
northern slopes of the Canary Islands, where, at middle eleva-
Downloaded from https://academic.oup.com/treephys/article/21/14/1039/1634463 by guest on 22 December 2023
tions, laurel forest has the characteristic appearance of a cloud
forest (Hollermann 1981).
The Canarian laurel forest is dominated by evergreen tree
species that are adapted to a humid Mediterranean climate
with moderate temperatures, where clouds mitigate the impact
of the drought periods characteristic of this climate type. Some
authors have related the Canarian laurel forest structurally
with the Mediterranean schlerophyllous vegetation (Goodall
1983) or with the tropical mountain forests (Hübl 1988, Lösch
and Fischer 1994), although, in terms of mineral nutrient com-
position, it is more similar to tropical forests (Köhl et al.
1996).
Gas exchange studies have been carried out on trees grow-
ing in a typical Mediterranean climate (e.g., Tenhunen et al.
1981, Pereira et al. 1986), in tropical and subtropical forests
(e.g., Zotz et al. 1995, Ishida et al. 1996, 1999, Eschenbach et
al. 1998) and in other types of forests (see review by Larcher
(1995)). However, little is known about the gas exchange
characteristics of trees in the Canarian laurel forest under natu-
ral conditions.
Previously, we characterized the stand structure and leaf
area distribution of a laurel forest stand in which Laurus
azorica (Seub.) Franco was the most common tree species
(Morales et al. 1996a, 1996b). Leaf characteristics along a
vertical canopy profile reflected the high leaf area index (LAI,
7.8) of the stand (Morales et al. 1996a) and the severe attenua-
tion of light by the canopy, which transmits only 2% of inci-
dent photosynthetically active radiation to ground level
(Aschan et al. 1994, 1997). In particular, we found that
L. azorica has a relatively high leaf plasticity index defined as
the difference between sun- and shade-leaf thickness relative
to sun leaf thickness (Carpenter and Smith 1981), indicating
that leaves of this species are highly shade-tolerant (Morales et
al. 1996c). It has also been shown that the ability of the
photosynthetic apparatus to withstand dehydration is related
to the vertical canopy profile (Jiménez et al. 1999).
The present study investigated leaf gas exchange character-
istics of L. azorica measured in the field in relation to canopy
position (upper and lower canopy) under a wide range of envi-
ronmental conditions occurring over the course of an entire
year.
1040 GONZÁLEZ-RODRÍGUEZ, MORALES AND JIMÉNEZ

Materials and methods
The study was conducted in a natural laurel forest stand lo-
cated on Agua García Mountain in Tenerife, Canary Islands
(28°27′32″ N, 16°24′20″ W; altitude 820 m). The site, on a 12°
NNE slope, has a humid Mediterranean climate with a 30-year
mean annual temperature of 14.0 °C (absolute maximum and
minimum of 39.0 and 0.2 °C, respectively), mean relative hu-
midity of 80%, and mean annual precipitation of 733 mm. Al-
though fog precipitation does not contribute significantly to
total throughfall (Aboal 1998), relative humidity, which is
about 30 °C in summer and 9 °C in winter. Mean annual tem-
perature during the study year was 14.4 °C. On most days, rel-
ative humidity exceeded 90%. High air vapor pressure deficits
(VPD) occurred only when there were winds from the Sahara
(Figure 1). Annual gross precipitation was 396.5 mm, most of
which occurred during autumn and spring, although light rain
occurred throughout the year. Thus, the climate at the study
site is mild without marked seasonality.
Diurnal courses of gas exchange
Diurnal courses of gas exchange differed according to the pre-

Downloaded from https://academic.oup.com/treephys/article/21/14/1039/1634463 by guest on 22 December 2023

above 80% on most days, mitigates the effects of drought.
The experimental stand comprises a mixed hardwood forest vailing environmental conditions. Figure 2 shows the diurnal
with six major tree species (L. azorica, Persea indica (L.) courses of gas exchange in sun leaves during 2 days with con-
K. Spreng (both Lauraceae), Myrica faya Ait. (Myricaceae), trasting environmental conditions. On a cloudless day (Janu-
Erica arborea L. (Ericaceae), Ilex canariensis Poir. and Ilex ary 25, 1995) when VPD was high (> 1.5 kPa), Amax was only
perado Ait. subsp. platyphylla (Webb and Berth.) Tutin (both 8 µmol m –2 s –1, because it was limited by low gs and the result-
Aquifoliaceae)). We studied adult trees of L. azorica, the most ing decrease in Ci. In contrast, during the wet conditions of
representative species of the laurel forest and the most abun- March 29, 1995, when VPD was low (< 0.5 kPa), Amax reached
dant species in our experimental stand (38%). A detailed de- 11.6 µmol m –2 s –1 and high values of E and gs were recorded
scription of the site and stand structure has been presented by (8 and 650 mmol m –2 s –1, respectively). On both days, A paral-
Morales et al. (1996a, 1996b). Leaf area index is 7.8 (Morales leled the diurnal pattern of photosynthetic photon flux density
et al. 1996a), which is similar to the LAI of Japanese, ever- (PPFD).
green broad-leaf forests (Kawanabe 1977, Tadaki 1977), low- The CO2 concentration of ambient air (Ca ) in the upper can-
land rainforest in Malaysia (Kato et al. 1974, Yoda 1974) and opy varied from 345 µmol mol –1 in the morning to 340 µmol
some evergreen conifer forests (Waring 1983).
A wooden scaffolding tower was erected at the site to access
leaves of L. azorica at the top (sun leaves, 18 m above ground)
and in the lower part (shade leaves, 12 m above ground) of the
canopy. Morales et al. (1996c) determined that total chloro-
phyll concentration per leaf dry mass and specific leaf area
(SLA) of sun and shade leaves were 4 g kg –1 and 66.7 cm 2 g –1
and 5.1 g kg –1 and 111.1 cm 2 g –1, respectively.
Climatic conditions during the study were recorded by a
meteorological station installed at the top of the tower (data
logger, with temperature and humidity sensors, Delta-T, Cam-
bridge, U.K.; quantum sensor LI-190 SA, Li-Cor, Lincoln,
NE; and rain gauge, Skye Institute, Llandrindal, U.K.).
Carbon assimilation rate (A), internal CO2 concentration
(Ci ), leaf conductance (gs ) and transpiration (E) were mea-
sured on fully developed current-year leaves with a portable
gas analyzer (LCA2, Analytical Development Company Ltd.,
Hoddesdon, U.K.) from the early morning to the evening. Care
was taken not to change the natural exposure of leaves during
gas exchange measurements. Measurements were made every
2 weeks from July 1994 to June 1995, when climatic condi-
tions were favorable. To obtain the response of shade leaves to
light, attached branches were exposed to direct sunlight and
light curves were made with the help of different layers of
white cloth.
Linear regression was used to explore relationships between
different parameters and correlation coefficients were calcu-
lated (r).

Results Figure 1. Mean daily air temperature (Tmean), air relative humidity
(RH) and air vapor pressure deficit (VPD) during the study period
Climatic conditions
(July 1994 to June 1995). Measurements were recorded 2 m above the
During the study, mean daily air temperature varied between canopy.

TREE PHYSIOLOGY VOLUME 21, 2001

 GAS EXCHANGE CHARACTERISTICS OF LAURUS AZORICA
Figure 2. Diurnal courses of photosynthetic photon flux density
(PPFD) at the leaf surface, leaf temperature (Tleaf), air vapor pressure
deficit (VPD), carbon assimilation rate (A), transpiration rate (E),
stomatal conductance (gs), leaf internal CO2 concentration (Ci ), in-
stantaneous water-use efficiency (WUE = A/E) and intrinsic wa-
ter-use efficiency (A/gs) measured in Laurus azorica leaves at the top
of the canopy (sun leaves) around the sampling tower on January 25,
1995 (a day with high VPD) and on March 29, 1995 (a day with low
VPD). Each value represents an individual measurement. For each
measurement day, the top panel shows the diurnal patterns of Tleaf
(䊊), VPD (䊉) and PPFD (solid line).
mol –1 at midday on January 25, 1995, and the corresponding
values for March 29, 1995 were 350 and 325 µmol mol –1. At
midday, mean CO2 concentration of the leaf intercellular
space (Ci ) and mean Ci /Ca ratio were 225 µmol mol –1 and 0.67
on January 25, 1995 compared with 260 µmol mol –1 and 0.77
on March 29, 1995, respectively.
A low instantaneous water-use efficiency (WUE = A/E) of
about 1.5 µmol mmol –1 was recorded at midday on both Janu-
ary 25 and March 29, 1995. Compared with instantaneous
WUE, intrinsic water-use efficiency (A/gs) was slightly higher
on the cloudless day with high VPD (January 25, 1995) mainly
because of the decrease in gs, but even on this day we found no
evidence of midday stomatal closure. The stomata remained
TREE PHYSIOLOGY ONLINE at http://heronpublishing.com
1041
open enough throughout the day to maintain relatively high
values of A and E (Figure 2).
Variation in gas exchange characteristics among canopy
leaves
In general, irradiance in the lower canopy was less than
200 µmol m –2 s –1 during the study period. There was a large
difference in A between sun and shade leaves, such that shade
leaves were near their light compensation point most of the
time. On any given day, maximum values of A (2 µmol m –2
Downloaded from https://academic.oup.com/treephys/article/21/14/1039/1634463 by guest on 22 December 2023
s –1), gs (130 mmol m –2 s –1) and E (2.5 mmol m –2 s –1) of shade
leaves represented about 25, 47 and 50%, respectively, of the
corresponding values of sun leaves (Figure 3). Instantaneous
and intrinsic WUE were also much lower in shade leaves than
in sun leaves, attaining maximum values of 0.8 and 14 µmol
mol –1, respectively, indicating that shade leaves had higher Ci
than sun leaves.
Analysis of total data
For data obtained at the top of the canopy (n = 550), there were
high correlations between light and leaf temperature r = 0.83),
carbon assimilation rate and stomatal conductance r = 0.47),
and carbon assimilation rate and transpiration r = 0.72). We
used boundary-line analysis using all of the data of each pa-
rameter plotted against every variable, to estimate the maxi-
mal parameter values when other variables were not limiting.
In sun leaves, the relationship between A and leaf temperature
indicated that the optimum temperature ranged between
20 and 26 °C. A plot of A versus photosynthetic photon flux
density (PPFD) yielded an Amax of 10.5 µmol m –2 s –1 and indi-
cated that A reached light saturation above 750 µmol m –2 s –1.
The plot of A versus gs did not pass through the origin because
the stomata were partly open when A was zero, and the rela-
tionship did not show a clear saturation curve. Values of gs
above 300 mmol m –2 s –1 were considered to be non-limiting
for sun leaves (Figure 4), whereas for shade leaves a gs value
of 125 mmol m –2 s –1 was considered to be non-limiting. In
shade leaves, Amax was about 5 µmol m –2 s –1 and A reached
light saturation above 500 µmol m –2 s –1 (Figure 5).
Relationships between gs and leaf temperature, PPFD and
VPD are shown in Figure 6. Stomatal conductance showed a
temperature response similar to that of A, although the optimal
range of temperatures was greater for gs than for A (15 to
26 °C). Maximum values of gs were even exhibited at low ir-
radiances (14 µmol m –2 s –1). Although gs decreased with in-
creasing VPD, relatively high gs values, near 300 mmol m –2
s –1, were maintained even at high VPDs, provided that light
and temperature were optimal.
Discussion
Although CO2 concentration, temperature and humidity dif-
fered with canopy height, the greatest vertical gradient was in
solar irradiance (Jarvis et al. 1976). The lower A in lower-can-
opy leaves compared with upper-canopy leaves reflected the
high light attenuation in this forest (Aschan et al. 1994, 1997),
1042 GONZÁLEZ-RODRÍGUEZ, MORALES AND JIMÉNEZ
Figure 3. Diurnal courses of photosynthetic photon flux density
(PPFD) at the leaf surface, leaf temperature (Tleaf), air vapor pressure
deficit (VPD), carbon assimilation rate (A), transpiration rate (E),
stomatal conductance (gs), leaf internal CO2 concentration (Ci), in-
stantaneous water-use efficiency (WUE = A/E) and intrinsic water-
use efficiency (A/gs) measured in Laurus azorica leaves in the upper
canopy (sun leaves) and lower canopy (shade leaves). Each value rep-
resents an individual measurement. For each leaf type, the top panel
shows the diurnal patterns of Tleaf (䊊) and PPFD (solid line).
which also resulted in leaves differing in morphology includ-
ing leaf area and thickness (Morales et al. 1996c). Estimated
values of Amax, based on light responses, were consistent with
expected differences between sun and shade leaves (Amax for
shade leaves was about 50% of that for sun leaves), and were
comparable with published values for C3 trees (Larcher 1995,
Lüttge 1997). Sun leaves of most tree species become light
saturated at 25–30% of full sunlight (Sprugel 1989). Sun and
shade leaves of L. azorica became light saturated at 37.5 and
25% of full sunlight, respectively. Our results indicate that
TREE PHYSIOLOGY VOLUME 21, 2001
Downloaded from https://academic.oup.com/treephys/article/21/14/1039/1634463 by guest on 22 December 2023
Figure 4. Relationships between carbon assimilation rate (A) and leaf
temperature (Tleaf), photosynthetic photon flux density (PPFD) at the
leaf surface, and stomatal conductance of water vapor (gs) in sun
leaves of Laurus azorica. Each value represents an individual mea-
surement.
L. azorica leaves are highly plastic, enabling them to adjust
their morphology and physiology to the microclimatic condi-
tions. We did not determine how the shade leaves respond to
short periods of direct irradiance (sunflecks). The duration and
quantity of exposure to sunflecks can greatly influence growth
(Pearcy 1988); i.e, up to 60% of the carbon gain of understory
plants is attributed to utilization of sunflecks (Pearcy et al.
1994).
Daily courses of gas exchange showed a typical response of
photosynthesis to light. During the morning as PPFD in-
creased, A, E and gs increased, whereas Ci decreased. On days
with low VPD, these parameters were mainly controlled by
PPFD; however, on days with high VPD, A and E decreased
because of a gradual reduction in gs throughout the whole day.
Although midday depression of gs as a mechanism for con-
serving water has been reported for temperate-zone plants
(Hodges 1967, Dougherty and Hinckley 1981), Mediterra-
nean-climate plants (Tenhunen et al. 1981, Pereira et al. 1986)
and tropical trees (Medina et al. 1978, Zotz et al. 1995, Pathre
et al. 1998, Ishida et al. 1999), we did not observe midday
stomatal closure in L. azorica.
 GAS EXCHANGE CHARACTERISTICS OF LAURUS AZORICA
Figure 5. Relationships between carbon assimilation rate (A) and
photosynthetic photon flux density (PPFD) at the leaf surface, and
stomatal conductance of water vapor (gs) in shade leaves of Laurus
azorica. Each value represents an individual measurement.
Compared with other species, L. azorica leaf photosyn-
thesis was relatively low; however, leaf photosynthesis in tree
species varies widely, ranging from about 3 to 30 µmol m –2 s –1
(Ceulemans and Saugier 1991). Generally, photosynthetic
rates of tropical rain forest trees are higher (10–16 µmol m –2
s –1) than those of broad-leaved evergreen trees of the subtrop-
ics and warm-temperate regions (6–12 µmol m –2 s –1) (Larcher
1995). The maximum value of A measured in L. azorica
(11.6 µmol m –2 s –1) and that obtained from a plot of all A val-
ues versus PPFD (10.5 µmol m –2 s –1) were within the limits of
both ranges. Photosynthetic rates in evergreen leaves are cor-
related with the potential for year-round photosynthesis and
leaf longevity (Schulze 1982, Reich et al. 1991, 1992, Kita-
jima et al. 1997). Leaf longevity, which is more than 3 years
for L. azorica, could account for its low photosynthetic rates.
During the study, the Ci /Ca ratio varied from 0.95 to 0.67,
which was higher than the range found in the Australian ever-
green species, Citrus L. and Eucalyptus L’Hér (Lloyd et al.
1992, Sheriff 1992), and a Mediterranean evergreen sclero-
phyllous shrub, Arbutus unedo L. (Beyschlag et al. 1987), sug-
gesting a relatively small stomatal limitation of A in
L. azorica. Relatively high Ci /Ca ratios have also been found
in several tropical rain forest species (Lloyd and Farquhar
1994, Ishida et al. 1996).
Stomatal conductance was sensitive to increasing VPD over
the range 0 to 3 kPa, decreasing exponentially but only until
the relatively high value of 300 mmol m –2 s –1 was reached, in-
dicating that little stomatal closure occurred, provided that
temperature and light were optimal. Only the low irradiances
in the early morning and late evening were not saturating for
TREE PHYSIOLOGY ONLINE at http://heronpublishing.com
1043
Downloaded from https://academic.oup.com/treephys/article/21/14/1039/1634463 by guest on 22 December 2023
Figure 6. Relationships between stomatal conductance of water vapor
(gs) and leaf temperature (Tleaf ), photosynthetic photon flux density
(PPFD) at the leaf surface and air vapor pressure deficit (VPD) in sun
leaves of Laurus azorica. Each value represents an individual mea-
surement.
the maximum value of gs. During the study, leaf temperature
oscillated between 12 and 35 °C, and temperatures below
15 °C and above 26 °C decreased gs. On some occasions, mea-
sured gs was lower than the corresponding value on the expo-
nential fitted curve of gs versus VPD. Maximum values of gs in
L. azorica were relatively high compared with those of woody
plants and trees of temperate forests, and more similar to those
of humid tropical forests than Mediterranean evergreen trees
(Körner 1995).
In general, L. azorica had high E (> 8 mmol m –2 s –1) and
low instantaneous WUE (mean = 1.5 µmol mmol –1), and also
low WUE when expressed as the ratio A/gs, which removes the
VPD effect (mean = 22 µmol mol –1). Similarly, Huc et al.
(1994) found that intrinsic WUE remained low throughout the
year in tropical rain forest tree species. However, in some
Mediterranean shrubs and trees, intrinsic WUE increases
markedly during the summer (Peñuelas et al. 1998, Flexas et
al. 2001). Low stomatal control in L. azorica is consistent with
the results obtained by Zöhlen et al. (1995) with porometric
techniques and Jiménez et al. (1996) with sap flow measure-
ments in which high values of whole-tree transpiration were
1044 GONZÁLEZ-RODRÍGUEZ, MORALES AND JIMÉNEZ
found throughout the year.
Plants have evolved two contrasting mechanisms (conser-
vative and prodigal water use) for dealing with limited water
supply (Passioura 1982). High gs and Ci /Ca ratios and low
WUE and A indicate that water use by L. azorica can be classi-
fied as prodigal or non-conservative, which corresponds with
the hydrolabile strategy defined by Larcher (1995). Non-con-
servative water use is common among plant species that are
subjected to only mild or short-duration droughts (Fischer and
Turner 1978). According to Passioura (1982), the most impor-
tant physiological distinction between prodigal and conserva-
tive plants is that the former probably have a greater ability to
maintain turgor when faced with a limited supply of soil water.
Laurus azorica exhibits efficient osmotic adjustment in re-
sponse to drought (González-Rodríguez et al. 1999) unlike its
congener L. nobilis L., a Mediterranean species, which shows
a sensitive adjustment of transpiration (Lo Gullo and Salleo
1988), and other shrubs growing in Mediterranean-type cli-
mates in California and Chile that have to endure severe and
prolonged summer droughts (Morrow and Mooney 1974).
However, even within the same environment, species display
different mechanisms for dealing with limited water supply as
has been found for Mediterranean shrubs (Larcher 1995,
Munné-Bosch et al. 1999) and in the Canarian laurel forest
(González-Rodríguez et al. 1999). Similarly, Lösch (1993)
found different mechanisms of water use in a study of Medi-
terranean and Canarian laurel forest trees.
We conclude that L. azorica exhibits great leaf plasticity as
exemplified by large differences in gas exchange characteris-
tics between upper-canopy sun leaves and lower-canopy
shade leaves that are modulated by the strong light attenuation
in the stand. This species has high E, gs and Ci/Ca and low A
and instantaneous WUE and A/gs. It responds to high VPD by
decreasing gs, but only to values high enough to maintain rela-
tively high A and E throughout the year. All gas exchange
characteristics fit well with a non-conservative water-use
strategy, which is appropriate for plants that are only subjected
to droughts that are mild or of short duration. Thus, this spe-
cies differs from its congener, L. nobilis, and other species of
the Mediterranean region that have to endure summer
droughts that are severe or of long duration or both. Overall,
the species is more similar to subtropical plants of humid re-
gions than to sclerophyllous plants of regions with a Mediter-
ranean-type climate.
Acknowledgments
This research was supported by DGICYT (PB 94-0580), Spanish
Government and the “Consejería de Educación, Cultura y Deportes”
of the Canarian Government. We express our gratitude to the latter for
the concession of a grant to Agueda M. González-Rodríguez. Thanks
also to Neil Abrey for checking the English.
References
Aboal, J.R. 1998. Los flujos netos hidrológicos y químicos asociados
de un bosque de laurisilva de Tenerife. Thesis Doctoral, Univer-
sidad de La Laguna, 184 p.
TREE PHYSIOLOGY VOLUME 21, 2001
Aschan, G., M.S. Jiménez, D. Morales and R. Lösch. 1994. Aspectos
microclimáticos de un bosque de laurisilva en Tenerife. Vieraea
23:125–141.
Aschan, G., R. Lösch, M.S. Jiménez and D. Morales. 1997. Energie-
bilanz von Waldbeständen in nicht-idealem Gelände-Abschätzun-
gen auf der Grundlage von standörtlicher Klimaerfassung und
flankierenden Gaswechselmessungen am Beispiel eines Lorbeer-
waldbestandes auf Teneriffa. EcoSys (Suppl.) 20:145–160.
Beyschlag, W., O.L. Lange and J.D. Tenhunen. 1987. Diurnal pat-
terns of leaf internal CO2 partial pressure of the sclerophyll shrub
Arbutus unedo growing in Portugal. In Plant Response to Stress.
Functional Analysis in Mediterranean Ecosystems. Eds. J.D. Ten-
Downloaded from https://academic.oup.com/treephys/article/21/14/1039/1634463 by guest on 22 December 2023
hunen, F.M. Catarino, O.L. Lange and W.C. Oechel. Springer-
Verlag, Berlin, pp 355–368.
Carpenter, S.B. and N.D. Smith. 1981. A comparative study of leaf
thickness among southern Appalachian hardwoods. Can. J. Bot.
59:1393–1396.
Ceulemans, P.J. and B. Saugier. 1991. Photosynthesis. In Physiology
of Trees. Ed. A.S. Raghavendra. Wiley-Interscience, New York,
pp 21–50.
Dougherty, W.J.S. and T.M. Hinckley. 1981. The influence of a se-
vere drought on net photosynthesis of white oak (Quercus alba).
Can. J. Bot. 59:335–341.
Eschenbach, C., R. Glauner, M. Kleine and L. Kappen. 1998. Photo-
synthesis rates of selected tree species in lowland dipterocarp rain-
forest of Sabah, Malaysia. Trees 12:356–365.
Fischer, R.A. and N.C. Turner. 1978. Plant productivity in the arid
and semiarid zones. Annu. Rev. Plant Physiol. 29:277–317.
Flexas, J., J. Gulías, S. Jonasson, M. Mus and H. Medrano. 2001. Sea-
sonal patterns and control of gas exchange in local populations of
the Mediterranean evergreen shrub Pistacia lentiscus L. Acta
Oecol. 22:33–43.
Goodall, D.W. 1983. Temperate broad-leaved evergreen forests. In
Ecosystems of the World, 10. Ed. J.D. Ovington. Elsevier, Amster-
dam, 242 p.
González-Rodríguez, A.M., M.S. Jiménez, D. Morales, G. Aschan
and R. Lösch. 1999. Physiological responses of Laurus azorica and
Viburnum rigidum to drought stress: osmotic adjustment and tissue
elasticity. Phyton 39:251–263.
Hodges, J.D. 1967. Patterns of photosynthesis under natural environ-
mental conditions. Ecology 48:234–242.
Hübl, E. 1988. Lorbeerwälder und Hartlaubwälder (Ost-asien, Medi-
terraneis und Makaronesien). Düsseldorfer Geobot. Koll. 5:3–26.
Huc, R., A. Ferhi and J.M. Guehl. 1994. Pioneer and late stage tropi-
cal rainforest tree species (French Guiana) growing under common
conditions differ in leaf gas exchange regulation, carbon isotope
discrimination and leaf water potential. Oecologia 99:297–305.
Hollermann, P. 1981. Microenvironmental studies in the laurel forest
of the Canary Islands. Mt. Res. Dev. 1:193–207.
Ishida, A., T. Toma, Y. Matsumoto, S.K. Yap and Y. Maruyama.
1996. Diurnal changes in leaf gas exchange characteristics in the
uppermost canopy of a rain forest tree, Dryobalanops aromatica
Gaertn. f. Tree Physiol. 16:779–785.
Ishida, A., T. Toma and Marjenah. 1999. Limitation of leaf carbon
gain by stomatal and photochemical processes in the top canopy of
Macaranga conifera, a tropical pioneer tree. Tree Physiol. 19:
467–473.
Jarvis, P.G., G.B. James and J.J. Landsberg. 1976. Coniferous forest.
In Vegetation and the Atmosphere. Ed. J.L. Monteith. Academic
Press, London, pp 171–240.
Jiménez, M.S., J. Èermák, J. Kuèera and D. Morales. 1996. Laurel
forests in Tenerife, Canary Islands: the annual course of sap flow in
Laurus trees and stand. J. Hydrol. 183:307–321.
 GAS EXCHANGE CHARACTERISTICS OF LAURUS AZORICA
Jiménez, M.S., A.M. González-Rodríguez and D. Morales. 1999. Ef-
fect of dehydration on the photosynthetic apparatus of sun and
shade leaves of laurel forest trees. Z. Naturforsch. 54c:704–710.
Kato, R., Y. Tadaki and H. Ogawa. 1974. Plant biomass and growth
increment studies in Pasohforest. Malay. Nat. J. 30:211–224.
Kawanabe, S. 1977. A subtropical broad-leaved forest at Yona, Oki-
nawa. In Primary Productivity of Japanese Forests, Productivity of
Terrestrial Communities. Eds. T. Shidei and T. Kira. JIBP Synt.,
Vol. 16, Tokyo, pp 268–279.
Kitajima, K., S.S. Mulkey and S.J. Wright. 1997. Decline of photo-
synthetic capacity with leaf age in relation to leaf longevities for
five tropical canopy tree species. Am. J. Bot. 84:702–708.
Köhl, K., R. Lösch, A.M. González-Rodríguez, M.S. Jiménez and
D. Morales. 1996. Foliar cation contents of laurel forest trees on the
Canary Islands. Flora 191:303–311.
Körner, C. 1995. Leaf diffusive conductances in the major vegetation
types of the globe. In Ecophysiology of Photosynthesis. Eds. E.-D.
Schulze and M.M. Caldwell. Springer-Verlag, Heidelberg, pp
463–490.
Larcher, W. 1995. Physiological plant ecology, 3rd Edn. Springer-
Verlag, Berlin, 506 p.
Lloyd, J. and G.D. Farquhar. 1994. 13C discrimination during CO2 as-
similation by the terrestrial biosphere. Oecologia 99:201–215.
Lloyd, J., J.P. Syvertsen, P.E. Kriedemann and G.D. Farquhar. 1992.
Low conductances for CO2 diffusion from stomata to the sites of
carboxylation in leaves of woody species. Plant Cell Environ. 15:
873–899.
Lo Gullo, M.A. and S. Salleo. 1988. Different strategies of drought
resistance in three Mediterranean sclerophyllous trees growing in
the same environmental conditions. New Phytol. 108:267–276.
Lösch, R. 1993. Water relations of Canarian laurel forest trees. In Wa-
ter Transport in Plants under Climatic Stress. Eds. M. Borghetti,
J. Grace and A. Raschi. Cambridge Univ. Press, Cambridge, pp
243–246.
Lösch, R. and E. Fischer. 1994. Vikariierende heidebuschwälder und
ihre kontaktgesellschaften in Makaronesien und Zentral-afrika.
Phytocoenologia 24:695–720.
Lüttge, U. 1997. Physiological ecology of tropical plants. Springer-
Verlag, Berlin, 384 p.
Medina, E., M. Sobrado and R. Herrera. 1978. Significance of leaf
orientation for leaf temperature in an Amazonian schlerophyll veg-
etation. Radiat. Environ. Biophys. 15:131–140.
Morales, D., M.S. Jiménez, A.M. González-Rodríguez and J. Èer-
mák. 1996a. Laurel forest in Tenerife, Canary Islands: I. The site,
stand structure and stand leaf area distribution. Trees 11:34–40.
Morales, D., M.S. Jiménez, A.M. González-Rodríguez and J. Èer-
mák. 1996b. Laurel forest in Tenerife, Canary Islands: II. Leaf dis-
tribution patterns in individual trees. Trees 11:41–46.
Morales, D., A.M. González-Rodríguez, J. Èermák and M.S. Jimén-
ez. 1996c. Laurel forest in Tenerife, Canary Islands: the vertical
profiles of leaf characteristics. Phyton 36:251–263.
Morrow, P.A. and H.A. Mooney. 1974. Drought adaptations in two
Californian evergreen sclerophylls. Oecologia 15:205–222.
Munné-Bosch, S., S. Nogués and L. Alegre. 1999. Diurnal variations
of photosynthesis and dew absorption by leaves in two evergreen
shrubs growing in Mediterranean field conditions. New Phytol.
144:109–119.
TREE PHYSIOLOGY ONLINE at http://heronpublishing.com
1045
Passioura, J.B. 1982. Water in the soil–plant–atmosphere continuum.
In Physiological Plant Ecology II. Water Relations and Carbon As-
similation. Encyclopedia of Plant Physiology, Vol. 12B. Eds. O.L.
Lange, P.S. Nobel, C.B. Osmond and H. Ziegler. Springer-Verlag,
Berlin, pp 5–33.
Pathre, U., A.K. Sinha, P.A. Shirke and P.V. Sane. 1998. Factors de-
termining the midday depression of photosynthesis in trees under
monsoon climate. Trees 12:472–481.
Pearcy, R.W. 1988. Photosynthetic utilization of lightflecks by under-
storey plants. Aust. J. Plant Physiol. 15:223–238.
Pearcy, R.W., R.L. Chazdon, L.J. Gross and K.A. Mott. 1994. Photo-
synthetic utilization of sunflecks: a temporally patchy resource on
Downloaded from https://academic.oup.com/treephys/article/21/14/1039/1634463 by guest on 22 December 2023
a time scale of seconds to minutes. In Exploitation of Environmen-
tal Heterogeneity by Plants: Ecophysiological Processes Above
and Below Ground. Eds. M.M. Caldwell and R.W. Pearcy. Aca-
demic Press, San Diego, pp 175–208.
Peñuelas, J., I. Filella, J. Llusiá, D. Siscart and J. Piñol. 1998. Com-
parative field study of spring and summer leaf gas exchange and
photobiology of the mediterranean trees Quercus ilex and Philly-
rea latifolia. J. Exp. Bot. 49:229–238.
Pereira, J.S., J.D. Tenhunen, O.L. Lange, W. Beyschlag, A. Meyer
and M.M. David. 1986. Seasonal and diurnal patterns in leaf gas
exchange of Eucalyptus globulus trees growing in Portugal. Can.
J. For. Res. 16:177–184.
Reich, P.B., C. Uhl, M.B. Walters and D.S. Ellsworth. 1991. Leaf
lifespan as a determinant of leaf structure and function among 23
Amazonian tree species. Oecologia 86:16–24.
Reich, P.B., M.B. Walters and D.S. Ellsworth. 1992. Leaf lifespan in
relation to leaf, plant, and stand characteristics among diverse eco-
systems. Ecol. Monogr. 62:365–392.
Schulze, E.-D. 1982. Plant life forms and their carbon, water and nu-
trient relations. In Encyclopedia of Plant Physiology. N.S. Physio-
logical Plant Ecology II, Vol. 12B. Eds. O.L. Lange, P.S. Nobel,
C.B. Osmond and H. Ziegler. Springer-Verlag, Berlin, pp
615–676.
Sheriff, D.W. 1992. Roles of carbon gain and allocation in growth at
different nitrogen nutrition in Eucalyptus camaldulesis and Euca-
lyptus globulus seedlings. Aust. J. Plant Physiol. 19:637–652.
Sprugel, D.G. 1989. The relation of evergreenness, crown architec-
ture, and leaf size. Am. Nat. 133:465–479.
Tadaki, Y. 1977. Forest biomass: Leaf biomass. In Primary Produc-
tivity of Japanese Forests, Productivity of Terrestrial Commu-
nities. Eds. T. Shidei and T. Kira. JIBP Synt., Vol. 16, Tokyo, pp
39–44.
Tenhunen, J.D., O.L. Lange and M. Braun. 1981. Midday stomatal
closure in Mediterranean type schlerophylls under stimulated habi-
tat conditions in an environmental chamber. II. Effect on the com-
plex of leaf temperature and air humidity on gas exchange of Arbu-
tus unedo and Quercus ilex. Oecologia 50:5–11.
Waring, R.H. 1983. Estimating forest growth and efficiency in rela-
tion to canopy leaf area. Adv. Ecol. Res. 13:327–354.
Yoda, K. 1974. Three-dimensional distribution of light intensity in a
tropical forest in West Malaysia. Jpn. J. Ecol. 24:247–254.
Zöhlen, A., A.M. González-Rodríguez, M.S. Jiménez, R. Lösch and
D. Morales. 1995. Transpiración y regulación estomática en
árboles de la laurisilva canaria medidos en primavera. Vieraea 24:
91–104.
Zotz, G., G. Harris, M. Königer and K. Winter. 1995. High rates of
photosynthesis in the tropical pioneer tree, Ficus insipida Willd.
Flora 190:265–272.