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UDK 630* 243 (001)




Matjaž ČATER , Primož SIMONČIČ

ABSTRACT: In view of evident changes in the reaction of European beech
(Fagus sylvaticaL.) to environmental changes, five plots with young trees of
the same age were established and studied on natural beech sites. Beech trees
were equally distributed along the light gradient and were divided according
to light conditions. The parameter used for evaluation of light conditions was
the indirect site factor (ISF) obtained by the WinScanopy analysis. Three
groups of canopy – light conditions were defined: stand conditions (ISF<20),
edge (20mature stand (ISF>25). In all categories light saturation curves and curves
describing dependence between intercellular CO2concentration in leaves and
assimilation rate (A-Ci) were measured under the same fixed parameters
(temperature, flow and CO2concentration, humidity, and light intensity) with
Li-6400, to compare responses between different light categories and different
plots within comparable light conditions.

Differences between canopy, edge and open area responses were confirmed
with high significance on all plots as well as between studied forest complexes.
On plots from Kočevje region, young beech indicated more shade tolerance, the
response to increased light intensity and different CO2concentration was greater
than the response of young beech on Pohorje plots within the same light intensities.
Responses of trees on plots in managed and virgin forest were also
different: young beech response in virgin forest plot was more shade-tolerant,
compared to response of young beech from plots in managed forest.

Key words: Beech, photosynthesis, light, CO, response


The more frequent and intensive pressures to which cies predominate; in particular natural beech forests
forests are exposed are connected with an increasing (Fagus sylvatica L.) (Kutnar 2003). The quality of
number of extreme events and consequently higher existing and future beech forests is closely connected
risk-rates of forest management decisions, especially on with our understanding of tree-response to different
marginaland extreme sites.The importance of autocht-light conditions, especially in an environment of reduhonous
tree species in preserving dynamic equilibrium ced light intensity under a mature canopy and in younger
and stability in forest ecosystems is frequently emphasi-development stages. Such knowledge leads to correct
zed (Zerbe 2002, Hannahet al. 1995, Stanturf and well-tuned spatial and temporal silvicultural measuandMadsen2002).
In Slovenia, where forests cover res, which may vary among different silvicultural syover
60 % of the country, sites of mixed broadleaf spe-stems. It is also directed at sustainable development and
a better future quality of forests (Kazda 1997). Solar


Dr. Matjaž Čater

radiation, temperatures and precipitation which inf


Dr. Primož Simončič

luence the distribution of plants are getting in times of

Slovenian Forestry Institute,Večna pot 2, 1000 Ljubljana,
intensive climatic extremes and climatic changes a new

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M. Čater, P. Simončič: PHOTOSYNTHETIC RESPONSE OF YOUNG BEECH (Fagus Sylvatica L.) ... Šumarski list br. 11–12, CXXXIII (2009), 569-576

dimension. Evident changes in distribution of plants and

species diversity consequently affect primary producti

vity (Callaghan etal. 2004).

Several key questions about the future response of
beech to expected changes such as temperature increase,
redistribution of precipitation and increase of atmospheric
CO concentration remain open and without answers.
Quotations in literature and research results are in most
cases unclear and sometimes even contradictory (Po or ter
1998,Lloyd andFarquhar 1996). Photosynthe
sis, the first estimate of net productivity could be
measured as the response of plants to different light intensity
or the response to different concentration of CO


which is entering the system in the controlled environment.
In spite of good understanding of processes of carbon
dynamics at leaf level in a changed CO


environment, it is difficult to make a prognosis of the fu

ture response of the whole plant, also because of the

short time interval of observations and numerous possible
interactions that have not yet been recognized.According
toBatič(2007) most changes by the increased
amount of atmospheric CO could be expected for C3


plants at the beginning of saturation curves, especially
for the plants that grow in reduced or minimal light conditions,
close to compensation point.Wemay therefore
expect most changes in shade tolerant species.

The research goal was to define range of photosynthetic
response in young beech in dependence of light
intensity and different concentration of CO between


three canopy conditions (shelter, forest edge and gap)
on different forest sites.

MATERIALAND METHODS – Materijali i metode

Research was performed on 10–15 year old beech
trees at five selected natural forest stands: at Kladje and
Brička in the Pohorje area, atVrhovo and the karstic-

Table 1 Research plots characteristics

Tablica 1.Značajkepokusnih ploha

dinaric area in Kočevski Rog –at Snežna jama (managed
forest) and Rajhenav (virgin forest) (Table 1).

air T
Prosječna godišnja
Tip tla
Growing stock
Drvna zaliha
Brička 1093 46o28’40’’ 15o15’40’ 1190 9,1 Dystric Cambisol 477
Kladje 1308 46o28’48’’ 15o23’24’’ 1066 9,2 Dystric Cambisol 390
Vrhovo 273 45o48’25’’ 15o18’11’’ 1138 9,4 Acric Luvisol 479
Sn. jama 875 45o39’15’’ 15o01’40’’ 1330 8,3 Rendzic Leptosol 612
Rajhenav 865 45o39’36’’ 15o03’36’’ 1330 8,3 Rendzic Leptosol 992

Both Brička and Kladje belong to the acidophilous
beech forest type Luzulo albidae-Fagetum (Urbančič
and Kutnar 2006) while Snežna jama and Rajhenav
belong to dinaric silver fir and beech forest type Omphalodo-
Fagetum (Kutnar andUrbančič2008).

At each location a research plot was established
100x100m in size, reaching from complete closure to
open sky conditions on all plots with little or no exposure.
The gradient of natural light conditions was obtained
by selecting young trees under a range of canopy
openness. On each fenced plot, for 24 young beech
trees in comparable light-intensity conditions, their potential
light environment was estimated with hemispherical
photos (Anonymous 2003). Fine tuning was
applied after pilot analysis, so that the light conditions
on all plots were comparable.The parameter used for
evaluation of light conditions was the indirect site factor
(ISF) (Wagner 1994), which is the relative proportion
of diffuse light intensity above a defined plant
compared to open/gap conditions, (without shading) in
percentage (%). Photos were taken with a digital Nikon
Coolpix 990 and calibrated fish-eye lens and analyzed
withWinScanopy software. In the process of hemispherical
photo analysis the vegetation period was defined
for each plot group separately; for the diffuse light
distribution a “Standard overcast sky” (SOC) model
was applied. For the calculation within the vegetation
period, the sun’s position was specified every ten (10)
minutes.The solar constant was defined as 1370W/m2,

0.6for atmospheric transmissivity and 0.15 for the proportion
of diffuse radiation compared to calculated direct
potential radiation.According to light conditions
three groups were defined: stand conditions (ISF<20),
edge (20the sheltering effect of a mature stand (ISF>25). Height
of trees on plots ranged from 40 - 70 cm under stand
conditions, from 70–110 cm under edge conditions and
from 110–220 cm in open area conditions. In each
group, four trees were randomly selected for measurement
of photosynthesis. In the same leaves nitrogen


concentration [mg/cm] was determined to compare

macronutrient status in different light categories (Leco

CNS-2000 analyzer) (Anonymous,2007).

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Light saturation curves were established to define
comparable ecophysiological response of net assimilation
(A) in beech leaves to different light intensities in
different plots and in comparable potential light conditions,
as described byPotočić etal. (2009).All photosynthesis
measurements were performed at a
constant temperature of the measurement block (20oC),
a CO concentration of 350 µmol/l, flow 500 µmol/s


and different light intensities: 0, 50, 250, 600 and


1200µmol/ms. Measurements started at ambient light
conditions that were reduced to reach zero, then followed
by a gradual increase toward maximum values, so
that stomata could adapt.

A-Ci curves were established to compare and define
assimilation response of trees (A) to different intercellular
CO concentrations (Ci): measurements were per


formed at constant light 600 µmol/ms, humidity,
constant block temperature 20 oC and flow 500µmol/s,
while ambient CO was varied as 0, 50, 100, 350, 700


and 1000 µmol/l. Maximal assimilation (A ) rates


and calculated compensation points (CP) for the light
saturation andA-Ci curves were used in comparisons
of trees between different plots. Both types of response
were measured with an LI-6400 portable system on at
least three sun leaves per plant, located in the upper
third of the tree-crown height on every plot. Twelve
trees were measured on each plot, four per same canopy
light conditions.

Water use efficiency of photosynthesis (WUE), a
quantitative measure of the instantaneous gas exchange
in leaves was expressed as the ratio of carbon gain per
water lost [mol HO/µmol CO] (Larcher 1995,


Lambersetal., 1998), while photosynthetic-use efficiency
(PNUE) as the carbon gain per unit leaf nitrogen
[µmol CO/gN] (Larcher 1995, Lambers et al.,


1998) for each light category, respectively.Atotal of 20
leaves were sampled per seedling in the upper crown
position, then cool-stored in airtight conditions. Fresh
leaves were weighed and scanned for the leaf area.
Leaves were dried at 105ofor 24 hours until constant
weight and weighed for the dry mass.

Analyses of variance (ANOVA) and post hoc LSD
analysis were used after testing data to meet conditions
of normality. Probability values of P<0.05 (*), P<0.01
(**) and P<0.001 (***) were considered significant.
Statistical data analysis was done with the programme

RESULTS – Rezultati istraživanja


The nitrogen content defined per leaf unit (mg/cm)
was different between Pohorje and Kočevje plots in canopy
(df ; F=105.13***), edge (df ; F=6.19*) and


gap conditions (df ; F=40.99***). On every plot, the


amount was highest in forest gap and lowest under
shelter conditions, except in virgin forest, with maximum
values at the forest edge (Table 2). Differences
between edge and open area conditions were not significantly
different on both plots from Kočevje (Table 3).

Table 2
Average leaf nitrogen content per leaf area, water use efficiency (WUE) and photosynthetic nitrogen use efficiency
(PNUE) on plots (means ± SE, n=8)

Tablica 2.Prosječni sadržaj dušika po jedinici površine lista, efikasnost uporabe vode (WUE) i fotosintetska efikasnost uporabe
dušika (PNUE) na plohama (sredine± SE, n=8)

Nitrogen (N) [mg/cm
] WUE [mol H
O/µmol CO
] PNUE [µmol CO
Brička 7.5±1.2 9.7±1.3 12.7±1.4 20.0±2.1 18.7±2.3 16.6±2.8 0.04±0.002 0.03±0.002 0.04±0.004
Kladje 8.8±1.0 12.4±1.7 18.2±1.9 22.9±2.4 20.4±1.5 15.8±2.2 0.04±0.005 0.03±0.003 0.02±0.003
Vrhovo 4.5±0.9 8.6±1.5 10.9±2.1 17.9±1.8 12.2±2.7 8.9±1.8 0.06±0.008 0.06±0.003 0.04±0.005
Sn. jama 4.7±0.8 10.4±0.7 10.7±1.6 19.9±3.3 18.8±3.2 15.2±1.9 0.10±0.006 0.08±0.002 0.08±0.003
Rajhenav 3.4±0.8 9.0±0.6 8.9± 1.3 20.3±2.6 20.9±3.1 19.9±3.3 0.08±0.006 0.06±0.002 0.04±0.006

The values for water use-efficiency (WUE) were
highest under shelter on all plots, ranging from 17.9

22.9mol HO/µmol CO, with the exception Rajhenav,


where maximum values were measured at the forest
edge (20.9 mol HO/µmol CO) (Table 2). Although the


value measured at the forest edge in Rajhenav was
greater from that measured in gap it was statistically
not significant (20.9 mol HO/µmol CO compared to


20.3mol HO/µmol CO, respectively).Asimilar rela


tion was determined for photosynthetic nitrogen use efficiency
(PNUE), highest under shelter at Snežna jama

(0.10µmol CO/gN).The highest values for the maxi


mum assimilation rate (A ) was measured in the open


(gap) atVrhovo, followed by the plots from the Pohorje
complex (Brička and Kladje), while the lowest values
were measured on plots in Kočevski Rog (Snežna
jama, Rajhenav) (Table 4).

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Table 3
Differences in leaf nitrogen between categories on
plots (AVAR and post hoc LSD
analysis): 1- shelter; 2 - edge; 3 - open light conditions;
NS… non-significant differences

Tablica 3.Razlike u sadržaju dušika u lišću izmedju različitih
kategorija na plohama (AVAR i post hoc LSD
analiza): 1- zastor; 2 - rub; 3 - otvoreno; NS… nesignifikantne

df (2, 21)
Ploha F p LSD
Brička 37.173 0.000 1-2 p=0.0021
1-3 p=0.0000
2-3 p=0.0001
Kladje 93.225 0.000 1-2 p=0.0000
1-3 p=0.0000
2-3 p=0.0000
Vrhovo 70.205 0.000 1-2 p=0.0000
1-3 p=0.0000
2-3 p=0.0001
Sn. Jama 68.914 0.000 1-2 p=0.0000
1-3 p=0.0000
2-3 p=0.7419 NS
Rajhenav 88.634 0.000 1-2 p=0.0000
1-3 p=0.0000
2-3 p=0.7419 NS

Differences between canopy, edge and open area responses
were confirmed with high significance on all
plots (Table 5) except in Rajhenav (virgin forest),
where no differences between canopy and edge area
conditions (df ; F=0.13; NS) were confirmed.


The response to maximum light in gap conditions
between Snežna jama and Rajhenav showed also no differences.
Assimilation responses to light were higher in
all categories in the virgin forest. The calculated light
compensation point (LCP) for edge and gap conditions
in Rajhenav and Snežna jama were practically the same


(20µmol/ms), values on other plots followed maximal
assimilation rates, respectively (data not shown).

The assimilation response of young beech between
the two forest complexes was also significantly different
between canopy (df ; F=285.99***), edge (df ;


F=171.68***) and gap conditions (df ; F=93.30***).


Table 4
Average values of maximum assimilation rates

(A ) (means ± SE, n=24)


Tablica 4.Prosječne vrijednosti maksimalne asimilacije
(Amax) (sredine ± SE, n=24)

Table 5
Differences in maximum assimilation rates (A )


on plots (AVAR and post hoc
LSD analysis): 1- shelter; 2 - edge; 3 - open light
conditions; NS… non significant differences

Tablica 5.Razlike u maksimalnoj asimilaciji (Amax) na plohama
(AVAR i post hoc LSD
analiza): 1- zastor; 2 - rub; 3 - otvoreno; NS… nesignifikantne

df (2, 21)
Ploha F p LSD
Brička 58.681 0.000 1-2p=0.0000
1-3 p=0.0000
2-3 p=0.0001
Kladje 28.804 0.000 1-2 p=0.0002
1-3 p=0.0000
2-3 p=0.0049
Vrhovo 442.675 0.000 1-2 p=0.0000
2-3 p=0.0000
Sn. Jama 75.266 0.000 1-2 p=0.0000
1-3 p=0.0000
2-3 p=0.0000
Rajhenav 12.495 0.000 1-2 p=0.6002 NS
2-3 p=0.0006

Maximum assimilation values forA-Ci curves (A


) (Table 6) showed similar reaction of trees as in the
case of maximum assimilation values measured for the
light curves (A ) (Table 4).


The response of young beech was greatest in the category
of canopy gap and lowest under shelter on all
plots.The highest response to increased CO concentra


tion was evident atVrhovo, followed by plots on Pohorje
(Brička and Kladje) and lowest in Kočevski Rog
(Rajhenav, Snežna jama). Comparison of the calculated
compensation point for CO between light catego


ries showed no significant differences (data not
shown). Differences between forest complexes were
statistically different in all categories: canopy (df ;


F=6.47**), edge (df ; F=13.17**) and gap conditions


(df ; F=33.41***).


Table 6
Maximum assimilation values for A-Ci curves
(A ); (means ± SE, n=24 trees).

max A-Ci

Tablica 6.Maksimalna asimilacija (Amax A-Ci);
(sredine ± SE, n=24 stabla).

Amax (µmol /m
Brička 7.3±0.4 9.8±0.8 11.9±1.1
Kladje 8.3±0.3 9.7±0.5 10.7±0.9
Vrhovo 6.1±0.4 9.3±0.4 13.2±0.6
Snežna jama 4.8±0.4 6.5±0.5 8.0±0.7
Rajhenav 7.1±0.3 7.2±0.5 8.2±0.6

(µmol /m
Brička 6.1±0.7 7.7±0.6 12.3±0.5
Kladje 8.8±0.6 9.6±0.5 10.4±0.6
Vrhovo 5.4±0.4 7.1±0.8 13.4±1.0
Snežna jama 3.4±0.4 5.4±0.4 9.4±0.6
Rajhenav 7.8±0.4 8.3±0.6 9.7±0.6

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In the Kočevski Rog area, differences among the tree ged one (at Snezna jama).Assimilation rates were higlight
categories are smaller in the virgin forest where no hest in all categories in virgin forests, despite the comdifferences
in response between canopy and edge were parable amount of nitrogen in leaves on the two plots.
confirmed (Post hoc LSD, p=0.0969) than in the mana-


Sensitivity of photosynthesis is similar for all C


plants and is in proportion with mesophyll CO concen


tration (Farquhar etal. 1980). Many studies indicate
that trees have higher mesophyll resistance for CO,


consequently lower photosynthesis and are therefore
more susceptible to the increase of atmospheric CO


concentration. In the view of climatic changes numerous
and often contradictory conclusions are being presented
about the response of plants and future development of
tree adaptation to environmental changes, especially due
to temperature increase (Körner, 2006), decrease in
amount of precipitation and changes in water supply
(Davies 2006) and increase of atmospheric CO con


centration (Ziska andBunce2006).
Light, nutrients, water and CO are abiotic parame


ters, necessary for the plant growth. Efficiency and
photosynthetic regulation are governed by ribulose-1,5
biphosphat carboxylase (rubisco), which is genetically
defined (Cheng etal. 1998). In general, by higher atmospheric
CO; protein synthesis in leaves increases,


stomatal aperture decreases, water use-efficiency and
C/N relation on the leaf level are increased, while on
the whole plant level growth is stimulated (Kimball
1993, Ghannoum et al. 2000).

By the increased amount of CO photosynthesis per


unit of the leaf area would increase, which is dependent
on the nitrogen supply. Respiration and root activities
would also increase, while biomass would be allocated
into roots (sweet chestnut) or increased proportionally
over whole plant (beech), which indicates a species-
specific response (Kohen etal. 1993).

In spite of relatively good insight into processes of
carbon dynamics on the leaf level in changed CO en


vironment it is difficult to make a prognosis of future
response of the whole plant also because of a short-
time interval of observations and numerous possible interactions
that haven’t been recognized yet (Increased
WUE might stimulate development of foliar fungi
(Thompson andDrake1994) while more sugars in
assimilation apparatus might stimulate the development
of pathogens and infections (Hibberd et al.
1996) etc.). Recent research quote up to 30 % increase
of growth in ambient with two times higher CO envi


ronment (Medylin etal. 2001).Asmaller probability
that such increase would reflect in long term growth in

assimilation was confirmed by Batič 2007, where

growth only increased at the beginning, and was later

reduced in time. Our analysis conirmed the differences
in response between beech under shelter, at the forest
edge and in the open. In spite of these differences, the
highest assimilation rates were measured on the research
plots of Pohorje complex and lowest on the
plots of Kočevski Rog. Results also indicate a different
response of young beech between managed forest
(Snežna jama) and virgin forest (Rajhenav); differences
between light categories were more pronounced
in the managed forest, while in virgin forest response to
same light conditions was more intense than in managed
forest. Photosynthetic yield in all categories was
higher in virgin forest. Light compensation point was
higher on plots of Pohorje complex compared to plots
in Kočevski Rog (data not shown).

Water use efficiency (WUE) was in all cases highest
under shelter and lowest in open conditions, similar to
photosynthetic nitrogen use-efficiency (PNUE).

Self shading and nitrogen redistribution within
whole plant could potentially underlie the degree of
photosynthetic acclimation to elevated CO (Takeu


chi et al. 2001); it is clear that interactions with other
potential environmental variables (light, nutrients) will
determine the regulation of carbon sources and sinks at
the leaf level (Lewis etal. 2002), but the ability to utilize
the knowledge and predict a whole plant response is
still limited and subject of controversy (Poorter 1998,

There were no significant differences in the content
of leaf nitrogen between plots. Leaf nitrogen values
were on all plots (expressed in units per leaf area) highest
in open conditions, without shading. Response of
young beech to different CO concentrations was simi


lar to response of young beech to different light intensity;
differences between managed and virgin forest
were even bigger under canopy and edge conditions.

In Kočevski Rog young beech is more shade tolerant,
relative response to increased light intensity and different
CO concentration is higher than response of young


beech in Pohorje within same light intensities. Responses
in managed and virgin forest are different: in the virgin
forest young beech trees are more shade-tolerant,
reaction of different light categories to elevated CO
concentration is similar and more homogenous, compared
to managed forest where differences between cate

gories are more pronounced. Kočevje region (Snežna

jama and Rajhenav) is well known for its forest manage

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M. Čater, P. Simončič: PHOTOSYNTHETIC RESPONSE OF YOUNG BEECH (Fagus Sylvatica L.) ... Šumarski list br. 11–12, CXXXIII (2009), 569-576

ment, with a long tradition of a sustainable and close-tonature
approach, with a single-tree selection method
(Diaci 2006). In contrast, in the Pohorje complex, with
potential beech sites, Norway spruce has been favored in
the last century and nowadays beech is gradually replacing
spruce either by underplanting or by natural regeneration
(Diaci 2006). In case of the response in
young beech trees, differences in assimilation rate may
reflect not only the different forest management history,
but also a different genetic background.


Autors wish to thank dr. Nenad Potočić, from Šumarski
Institut Jastrebarsko, Croatia, forconstructive
suggestions on the manuscript text and revision of titles
and abstract in Croatian language.The article is a result
of the “L4-6232; Carbon dynamic in natural beech forest”
project, financed by the Slovenian Research
Agency and Programme research group.

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čimbenik (ISF), dobiven analizom hemisfernih snimaka pomoću sustava Win-
Scanopy: zastor krošanja (ISF<20), rub sastojine (20bez zastora krošanja (ISF>25), koji su bili jednaki na svim plohama. Za izmjere
fotosintetskog kapaciteta, krivulje svjetlosnog zasićenja (0, 50, 250, 600
i 1200 µmol/m2s) i A-Ci krivulje (0, 100, 400, 700 i 1000 µmol CO2/l)
dobivene su pomoću Li-Cor LI-6400 u kontroliranom okruženju (temperatura,
protok i koncentracija CO2, zračna vlaga). Analize sadržaja dušika u lišću napravljene
su Leco CNS-2000 analizatorom.

Potvrđene su signifikantne razlike u reakciji mladih bukava između odabranih
kategorija, kao i između različitih šumskih kompleksa. Mlade bukve na plohama
iz Kočevskog pokazale su veću toleranciju na sjenu, a odziv na porast
koncentracije CO2je pri istim intenzitetima osvijetljenosti bio veći nego kod
mladih bukava iz Pohorskog kompleksa. Odziv mladih bukava bio je sig ni fi kant
no različit između prašume (Rajhenav) i gospodarske šume unutar istog
šumskog kompleksa: odziv u prašumi pokazuje veću toleranciju na sjenu.

Ključne riječi:Bukva, fotosinteza, svjetlo, CO2, odziv

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SAŽETAK: Glede uočenih promjena u reakciji bukve (Fagus sylvaticaL.) u
odnosu na ekološke promjene, odabrano je pet ploha mlade bukve jednake
starosti na prirodnim staništima, koje su bile jednakomjerno raspoređene na
svjetlosnom gradijentu od zastora odrasle sastojine, šumskog ruba do svjetlosnih
uvjeta na otvorenome. Kriterij za grupiranje bio je neizravni stanišni