South African Journal of Botany 132 (2020) 30
37

Contents lists available at ScienceDirect

South African Journal of Botany

journal homepage: www.elsevier.com/locate/sajb

Seasonal variation of fig tree (Ficus carica L.) physiological characteristics

reveals its adaptation performance
Aroua AMMARa,*, Imed BEN AISSAb, Messaoud MARSa, Mohamed GOUIAAc
a
Research Unit on Agrobiodiversity (UR13AGR05), Higher Agronomic Institute, Chott-Mariem, IRESA - University of Sousse, Tunisia
b
Research Unit on Integrated Horticultural Production (UR13AGR09), Regional Research Centre on Horticulture and Organic Agriculture, BP 57, 4042, Chott-Mar-
iem, IRESA - University of Sousse, Tunisia
c
Research Unit on Conservation and Valorization of Plant Resources (UR13AGR07), Higher Agronomic Institute, Chott-Mariem, IRESA - University of Sousse, Tunisia

A R T I C L E I N F O A B S T R A C T

Article History: In the context of climate change, understanding fig trees physiology is very important, since the areas of fig pro-
Received 20 September 2019 duction registered a large climate variability that severely affects fruit quality. The present study was conducted to
Revised 3 April 2020 evaluate the seasonal changes of physiological leaf behaviour of two Tunisian fig cultivars, Zidi and Bither Abiadh
Accepted 13 April 2020
during two years. Plants were subjected to full irrigation to maintain the soil moisture around pot capacity through
Available online 17 May 2020
the experimental period. Leaf gas exchange characteristics (net photosynthetic rate (PN), stomatal conductance
Edited by B Ncube (gs), and transpiration rate (E)), chlorophyll content index, leaf temperature and the maximum photochemical effi-
ciency of PSII (Fv/Fm) were measured. Results showed that the photosynthetic characteristics changed during the
Keywords:
growing season. Changes of photosynthetic CO2 assimilation, stomatal conductance, and chlorophyll content were
Ficus carica
not significantly affected by cultivar, and their variation was attributed to year. Seasonal changes of PN and gs
Adaptation
were similar suggesting that both processes are dependent. Indeed the best photosynthetic performance of fig
Photosynthesis
Stomatal conductance occurred about 80-87 days after bud break (spring) of the second season of growth, being supported by a high sto-
Chlorophyll matal aperture. The lowest values of PN, E and gs were recorded 156 days after bud break (summer) for 2017. The
Transpiration decrease of gas exchange rates and high temperatures were registered during the same dates. A little increase of
Tunisia the photosynthetic characteristics and chlorophyll content was noticed in the autumn period, about 180 days after
bud break, until a definitive decrease at the end of the growing season with the occurrence of low winter tempera-
ture. These results revealed particular adaptive behaviour and could be used as preliminary criteria to predict the
future impact of environmental abiotic stresses on ecophysiological fig tree behaviour.
© 2020 SAAB. Published by Elsevier B.V. All rights reserved.

1. Introduction
In arid and semi-arid regions of the Mediterranean basin, the cli-
mate is characterized by combined abiotic stresses such as low and
variable rainfall frequency, high sunlight intensity and elevated air
temperature (Giorgi and Lionello, 2008). A wide range of environ-
mental conditions may affect the photosynthesis activity as well as
morpho-physiological behaviour of perennial crops (leaf orientation
toward the radiation flux, anatomical leaf structure and morphology,
etc.) (Proietti and Famiani, 2002). For this reason, there is an increas-
ing interest in understanding trees physiology to gather the most
recent information about the effects of environmental factors and
their interactions with plants. Many research activities aim to predict
the future impact of increased aridity in these areas on the variation
of gas exchange in staple crops. Such studies help in deciding or
understanding the distribution and performance of certain fruit tree
species (Higgins et al., 1992).
* Corresponding author.
E-mail address: ammar_aroua@yahoo.fr (A. AMMAR).
In this regard, fig (Ficus carica L.), despite being widely cultivated in
the Mediterranean basin, few studies encompass data from its physio-
logical behaviour under similar and/or different climatic and edaphic
conditions (Higgins et al., 1992; Can et al., 2000; Kutlu et al., 2000; Can
and Aksoy, 2007). Hence, it is important to understand the physiologi-
cal characteristics of the fig tree to improve horticultural practices like
irrigation, fertilization, caprification, pruning, etc. Measuring gas
exchange capacity of the fig is necessary to evaluate its ability to grow
in different environmental conditions and to overcome low yields (Can
et al., 2008). Although few studies have been made on the fig resilience
to environmental stresses, previous studies showed that the fig photo-
synthesis rate (PN) was strongly affected by the genotype (Can et al.,
2000) and varied between clones of the same fig cultivar (Kutlu et al.,
2000). A study of seasonal changes in photosynthetic activity and sto-
matal conductance from April (leaf emergence) to August (fruit matura-
tion) showed significant changes during leaf development (Vemmos et
al., 2013). Can and Aksoy (2007) proved that PN was high in August due
to the increase of carbohydrate demand during fruit growth and matu-
ration. Gonza
lez-Rodríguez and Peters (2009) found that PN was
directly correlated with stomatal conductance, transpiration, leaf area

https://doi.org/10.1016/j.sajb.2020.04.020
0254-6299/© 2020 SAAB. Published by Elsevier B.V. All rights reserved.
 A. AMMAR et al. / South African Journal of Botany 132 (2020) 30
37
and relative chlorophyll content. They hypothesized that fig could be
considered as ‘delayed greening species’, since it showed pick PN and
chlorophyll content after full leaf expansion, which could be a strategy
to cope with environmental stresses and growth conditions.
The present study aims to investigate the physiological behaviour
over two seasons of two fig cultivars (Zidi and Bither Abiadh) of com-
mercial interest in Tunisia.
2. Materials and methods
2.1. Experimental site and plant material
This study was conducted in the experimental station of the
Regional Research Centre on Horticulture and Organic Agriculture
(CRRHAB), Chott-Mariem, Sousse (35°550 16.4"N 10°340 08.1"E), situ-
ated in the semi-arid center-east of Tunisia.
Five-year-old plants of fig cultivars Zidi ‘ZD’ (Smyrna type) and
Bither Abiadh ‘BA’ (San Pedro type) were used. Eight plants from
each cultivar were grown in 50L pots containing a substrate mixture
of 50% peat, 25% perlite, 15% sand and 10% compost with a potential
retention capacity of 50% and a permanent wilting point of 12%.
Plants were protected from rainfall with a plastic rooftop. During the
experiment, all plants were well-watered to pot capacity. All meas-
urements were performed under similar climatic conditions and
when soil moisture level was high after full irrigation.
2.2. Gas exchange capacity and leaf temperature measurements
Gas exchange measurements were undertaken in summer and
autumn seasons from August to November, corresponding to 156 days
after bud break (DAB) to 235 DAB in 2017. In 2018, the experiment was
conducted in spring, summer and autumn seasons from April to Novem-
ber, corresponding to 80 DAB to 255 DAB, during days with sunny sky.
Net photosynthetic assimilation rate PN (mmol m
2 s
1), transpiration
rate E (mmol m
2 s
1) and leaf temperature Leaf T (°C) measurements
were performed using a portable gas exchange system (CI-340; CID, Inc.,
USA). Leaf stomatal conductance gs (mmol m
2 s
1) was measured using
a leaf Porometer (model SC-1 of Decagon). These measurements were
always done on the uppermost fully expanded leaves and 16 replicates
were measured approximately from 8:00 am to 12:00 am.
Fig. 1. Daily courses of mean (Tmean24h) and maximum air (Tmax24h) temperature (Ta) (°C) during the experimental period of 2017 and 2018. DOY: day of year.
31
2.3. Chlorophyll Fluorescence and chlorophyll content index
Leaf chlorophyll fluorescence and chlorophyll content index
measurements were carried out using Portable pulse modulated
chlorophyll fluorometer OS5P (ADC, Bioscientific) and CCM-200plus
(Opti-sciences, USA) respectively. The minimal (F0) and maximal
fluorescence (Fm) emissions were assessed in leaves after 30 min of
dark adaptation, using clips for leaves and the Maximum photochem-
ical efficiency of photosystem II was calculated as follows:
Fv=Fm ¼ ðFm
F0Þ=Fm
2.4. Microclimate monitoring
The air temperature was recorded during the experimental period
using a high accuracy humidity, temperature and dew Point data log-
ger (EL-USB2+ sensor, Lascar Electronics, Inc.).
2.5. Data analysis
Statistical analyses were performed using IBM SPSS Statistics for
Windows, version 21 (IBM Corp., Armonk, N.Y., USA). The comparison of
mean values was made by one-way analysis of variance (ANOVA), fol-
lowed by Duncan’s test as described by Bewick et al. (2004) and differen-
ces were considered significant at the significant level of P  0.05. A two-
way ANOVA was also performed with the GLM (general linear model)
procedure with cultivars as fixed effects and year as a random factor to
test the effects of year, cultivar and their interaction as described by Du
et al. (2019). The figures, the third order polynomial regression analysis
(with the correlation coefficients R2) used for depicting the relationships
between gs and diurnal temperature and between PN and diurnal tem-
perature were performed by Microsoft Excel (2007). XLSTAT software
2014 was used to determine the correlation between all the parameters.
3. Results and discussion
3.1. Environmental conditions
The environmental conditions at the experimental site (microclimate)
were characterized by temperature fluctuations in both seasons (Fig. 1).
32 A. AMMAR et al. / South African Journal of Botany 132 (2020) 30
37
Table 1.
Mean squares and effects of cultivar, year and their interaction on physiological parameters of fig tree leaves.
PN gs
Cultivar 0.00 ns 4648.693 ns
Year 709.768 ** 1119020.724 **
Cultivar x Year 231.446 ** 69278.765 **
ns and ** indicate non-significant or significant differences at P 0.01.
Air temperature (Ta) was high in summer 2017 between 180 and 244
DOY (days of the year: calendar days), when maximum values were
higher than 40 °C and mean air temperature was between 30 and 36 °C. A
higher Ta was also found in the summer of 2018, with the highest values
around 45 °C. During this period, the mean Ta was fluctuating between 25
and 35 °C. In the autumn period (from 245 to 335 DOY), a decrease in
mean Ta was noticed in both seasons reaching less than 25 °C. However,
pick temperatures were also observed during the autumn period of 2018.
3.2. Seasonal changes in leaf gas exchange capacity of fig cultivars
during the two growing seasons
3.2.1. Seasonal variation of stomatal conductance (gs) and leaf
temperature (Leaf T)
The evaluation of physiological traits of fig cultivars showed that
gs changed during the growing season, but there was no significant
cultivar effect. The only exceptions were observed in summer period
for both studied years. In fact, gs was affected by year and the interac-
tion between year and cultivar (Table 1).
As shown in Fig. 2, the highest gs values were recorded at the end of
the spring season (81
88 DAB of 2018) when temperatures were
between 28 and 31°C. The gs reached 435.4 and 370.6 mmol m
2 s
1 for
‘ZD’ and ‘BA’ respectively. The lowest values of gs were observed for both
cultivars about 156 DAB (summer) during 2017 when the diurnal tem-
perature reached more than 35°C. These trends confirm those reported
by Can et al. (2008) who found that fig cultivars were stressed in August
when high temperatures occurred, and gas exchange rates were signifi-
cantly low compared to those at June which marks the beginning of the
summer season. Other evidence has shown that stomatal limitation in
grapevines (Greer and Weedon, 2012) and apple trees (Greer, 2014) var-
ied progressively over the growing season with the changes of climatic
conditions particularly with the variation of air temperature.
Fig. 2. Seasonal variations of leaf stomatal conductance (gs) (§SD) measured in 2017 and 2018 for fig cultivars ‘ZD’ and ‘BA’. * indicates significant differences between cultivars at P
0.05. ** indicates significant differences between cultivars at P0.01.
E Leaf T Chlorophyll index Fv/Fm
3.795 ** 54.039ns 9.894 ns 0.04 **
0.00 ns 174.575** 7844.602 ** 0.314 **
4.173 ** 11.398 ns 0.577 ns 0.084 **
Short-Term recovery of gs was observed during the autumn period
(185
194 DAB of 2017 and 182
190 DAB of 2018) but did not reach
the values registered in 81-88 DAB (spring) of 2018. At the end of the
growing season, with the occurrence of signs of senescence in 2018
and the decrease of leaf temperature less than 25°C (Fig. 3), gs
decreased and reached a minimum of 75.9 and 102.4 mmol m
2 s
1for
‘ZD’ and ‘BA’ respectively (Fig. 2).
In another study leaf temperature was considered as the major
factor limiting the gas exchange capacity of fig trees grown under
rain-fed conditions (Can and Aksoy, 2007). In our case, mean values
of leaf temperature reached more than 30°C in summer period for
the two seasons (Fig. 3), while a decrease in leaf temperature values
was noticed by the end of the growing season which coincided with
the end of the annual vegetative cycle of fig plants.
3.2.2. Seasonal variation in photosynthesis (PN) and transpiration rates (E)
The interaction between year and cultivar was significant for PN and
E as shown in Table 1. The PN was positively correlated to E (r=0.553**)
(Table 2). As shown in Table 1, changes of PN was not affected by the
type of cultivar but was attributed to the effect of the year.
In the first year of experiments, some significant differences were
observed between the cultivars. Thus, PN of ‘ZD’ ranged between 4.6
and 9.5 mmol m
2 s
1, while in ‘BA’ PN was between 3.7 and
6.8 mmol m
2 s
1 (Fig. 4). The lowest values of PN and E (Figs. 4 and
5) were recorded in 156 DAB (summer) of 2017, as observed for gs,
associated with the elevated temperature reaching a maximum of
45.9 °C in 2017 (Fig. 1). PN and E were positively correlated to gs as
shown in Table 2, in agreement with previous findings for fig trees
(Gonza
lez-Rodríguez and Peters, 2009).
On the other hand, in the second year of experiments, both cultivars
showed a similar changing pattern of PN. The photosynthesis rate was
at maximum in 80
87 DAB (spring) and reached 12.4 and
 A. AMMAR et al. / South African Journal of Botany 132 (2020) 30
37 33

Fig. 3. Changes of leaf temperature (Leaf T) (§SD) measured in 2017 and 2018 for fig cultivars ‘ZD’ and ‘BA’. ** indicates significant differences between cultivars at P0.01.

Table 2.
Pearson correlation coefficients between leaf physiological parameters in fig cultivar.

Chlorophyll index gs Leaf T PN E Fv/Fm

Chlorophyll index 1
gs 0,101* 1
Leaf T 0,034 0,043 1
PN 0,045 0,393** 0,216** 1
E -0,116* 0,232** 0,218** 0,553** 1
Fv/Fm 0,267** 0,163** -0,279** 0,061 -0,007 1
*Significant at P 0.05 ** Significant at P 0.01.

Fig. 4. Seasonal variation of net photosynthesis (PN) (§SD) measured in 2017 and 2018 for fig cultivars ‘ZD’ and ‘BA’. * indicates significant differences between cultivars at P 0.05.
** indicates significant differences between cultivars at P0.01.

10.2 mmol m
2 s
1 for ‘BA’ and ‘ZD’ respectively. These results are simi- Clifford et al. (1997) in field-grown fig, and those in caprifigs (Pige et al.,
lar to values found by Vemmos et al. (2013) for ‘Mission’ and ‘Fracasana’ 2001) and in the fig ‘alma’ cultivar (Andersen, 1989). In contrast, PN val-
fig cultivars (12.9 mmol m
2 s
1), which are also comparable to those of ues were lower than those found in other fig cultivars (19.5, 31.2 mmol
34 A. AMMAR et al. / South African Journal of Botany 132 (2020) 30
37

Fig. 5. Seasonal variations of transpiration rate (E) (§SD) measured in 2017 and 2018 for fig cultivars ‘ZD’ and ‘BA’. * indicates significant differences between cultivars at P 0.05. **
indicates significant differences between cultivars at P0.01.

m
2 s
1) (Can et al., 2000) and clones of ‘Sarilop’ cultivar (Kutlu et al.,
2000).
At 87 DAB (spring of 2018), a significant difference was found
between the cultivars (Fig. 4), ‘BA’ had a higher photosynthetic activity
which may coincide with the first crop (breba figs) development (Psi-
misi et al., 2012). Minimum values of PN were recorded in 164 DAB
(summer) for both cultivars reaching 6.1 mmol m
2 s
1 for ‘ZD’ and
6.7 mmol m
2 s
1 for ‘BA’, then increased in 186 DAB corresponding to
autumn period with no significant differences associated to cultivars,
and finally decreased again in the end of the growing season reaching
1.2 mmol m
2 s
1 in ‘ZD’ and 1.5 mmol m
2 s
1 in ‘BA’ (Fig. 4).
The decrease of PN in mid-summer or winter period could be not
only due to stomatal limitations (r=0.393**) but also to non-stomatal
resistance mechanisms such as photo-inhibition due to high irradiance
conditions and the reduction of leaf area (Angelopoulos et al., 1996, Can
and Aksoy, 2007). Leaf temperature had also a great effect on rates of PN
as shown in Table 2. A positive correlation was found between PN and
leaf temperature. This result indicated that the photosynthesis rate was
rising with increasing leaf temperature. This may show that the reached
values of leaf temperature were optimum for photosynthetic mecha-
nism, and fig plants could have a good ability to adjust their photosyn-
thetic activity to their leaf temperature. Low values of PN in the summer
period could be also due to the consumption for vegetative growth and
reproductive processes as well (Proietti and Famiani, 2002).
As shown in Fig. 6, for both cultivars, maximum gs was found when
the diurnal temperature was around 30 °C and 32 °C. When tempera-
ture increases, viscosity of water declines and mesophyll conductance
increases. This mechanism improves the supply of water to evaporative
sites, increasing the turgor pressure of the guard cells, which enhances
consequently the stomatal aperture (Urban et al., 2017). Similarly,

Fig. 6. Variation of gs (mmol m
2 s
1) values of fig cultivars ‘ZD’ and ‘BA’ with the change of mean diurnal temperature (°C).
 A. AMMAR et al. / South African Journal of Botany 132 (2020) 30
37 35

Fig. 7. Variation of PN (mmol m
2s
1) values of fig cultivars ‘ZD’ and ‘BA’ with the change of mean diurnal temperature (°C).

maximum PN for ‘ZD’ was found when the diurnal temperature was
between 30 and 32 °C (R2=0.681), however for ‘BA’ PN reached the maxi-
mum level at 32 and 34 °C (R2=0.563) (Fig. 7). These temperatures were
higher than those mentioned as an optimum air temperature for potted,
glasshouse-grown fig (26 °C) in Higgins et al. (1992)’s report; and lower
than the temperatures recorded in Pisimisi et al. (2012)’s findings
(Maximum PN was found under higher air temperature between 34 and
38 °C). These differences could be related to the climatic conditions of
the studied area, to the age of plant and type of cultivar.
We noticed that transpiration rates tend to show high variability
among the season for both cultivars. Results revealed that the maximum
value of transpiration rates in 2017 was obtained for ‘ZD’ cultivar reach-
ing 2.19 mmol m
2 s
1 about 172 DAB (summer). While in 2018, maxi-
mum E value was recorded for ‘BA’ cultivar during the summer period
(95 DAB) and reached 2.5 mmol m
2 s
1 (Fig. 5). Can et al. (2008)
demonstrated that transpiration, in other fig cultivars, could reach more
than 10 mmol m
2 s
1 during June and August. These differences may
be associated to the fact that data reported by Can et al. (2008) was for
open field mature trees, which are naturally subjected to high daily var-
iations in terms of air temperature, humidity and light intensity, as E
was positively correlated to gs and leaf temperature (Table 2).
3.3. Changes in chlorophyll content index during the two growing seasons
Chlorophyll index was not significantly affected by cultivar and
the interaction between year and cultivar. However, it was affected
by the year (Table 1). In 2017, chlorophyll content values were rela-
tively low for both cultivars and did not exceed 15 units until the
end of the experimental period when it reached its minimum val-
ues (Fig. 8). During 2018, the changing pattern of chlorophyll

Fig. 8. Changes in chlorophyll content index (§SD) measured in 2017 and 2018 for fig cultivars ‘ZD’ and ‘BA’. * indicates significant differences between cultivars at P 0.05. ** indi-
cates significant differences between cultivars at P0.01.
36 A. AMMAR et al. / South African Journal of Botany 132 (2020) 30
37
content index was quite similar for both cultivars. Maximum values
were reached in 88 DAB (spring) and 193 DAB (autumn) for ‘ZD’
and ‘BA’ cultivars, respectively. A slight reduction of chlorophyll
content index was noticed up to 165 DAB (summer), followed by
an increase for both cultivars. Then, a decrease of chlorophyll con-
tent index was registered at the end of the growing season, reach-
ing 5.2 units. The change of leaf chlorophyll content could be
associated with the stages of leaf development. Vemmos et al.
(2013) reports that high values are related to full leaf expansion,
while a decrease of this parameter may be linked to leaf ageing and
appearance of signs of senescence (Guyer et al., 2014). Hailemi-
chael et al. (2016) added that the variability of chlorophyll content
may be generated by iron deficiency or another nutrient status dur-
ing the study.
3.4. Seasonal changes of the maximum quantum efficiency of the PSII
photochemistry (Fv/Fm)
In both studied years, the maximum photochemical efficiency of
PSII (Fv/Fm) changed with the growing season without significant
differences between cultivars (Fig. 9). The only exceptions were in
171 DAB (summer) in 2017 when cultivar ‘ZD’ had the Fv/Fm value
reaching 0.741, which is significantly higher than ‘BA’, which reached
0.600, and in 165 DAB (summer) and 185 DAB (autumn) in 2018,
when the Fv/Fm ratio was significantly high for ‘BA’ cultivar. Our
results showed a positive correlation between chlorophyll index and
the Fv/Fm ratio, and this parameter was affected significantly by cul-
tivar, year and interaction between them (Table 1). We infer that a
high level of photosynthetic pigments might enhance light absorp-
tion, consequently increasing the maximum yield of photosystem II.
In 155 DAB of 2017 (summer), the ratio Fv/Fm had dropped to a mini-
mum of 0.493 for ‘BA’ cultivar and to 0.640 for ‘ZD’; and at the end of
the growing season in 2018, it also decreased to less than 0.700. This
result could be enhanced by environmental stresses such as high-
intensity light and high temperatures. Also, the decrease of the Fv/Fm
ratio may be caused by the photo-inhibition of PSII or due to the
accumulation of reactive oxygen species which inhibits the repair of
photo-damaged PSII (Zhang et al., 2016). Furthermore, Palliotti et al.
Fig. 9. Seasonal variation of maximum photochemical efficiency of PSII (Fv/Fm) (§SD) measured in 2017 and 2018 for fig cultivars ‘ZD’ and ‘BA’. * indicates significant differences
between cultivars at P 0.05. ** indicates significant differences between cultivars at P0.01.
(2009) showed that year, cultivar and leaf position had no significant
effect on the Fv/Fm ratio and the decrease of Fv/Fm values was
induced by high temperatures during the summer season. High val-
ues of the Fv/Fm ratio may indicate the absence of photo-oxidative
damage in summer as a consequence of the activation of photopro-
tective mechanisms (Vitale et al., 2012). Indeed, the maximum quan-
tum yield of PSII, given as Fv/Fm ratio, is generally used as an
indicator to evaluate the performance of the photosynthetic appara-
tus (Mlinari
c et al., 2016).
4. Conclusion
The aim of this study was to evaluate the seasonal variation of
some physiological traits of two different fig tree types (Smyrna and
San Pedro) grown under conditions of optimal water availability.
The results revealed an adaptive behaviour of fig to cope with
stressful situations such as high temperatures. Indeed, temperature
affects fig physiological processes, including photosynthesis and tran-
spiration. Both, PN and E, are regulated by stomatal conductance (gs)
and they mutually affect each other to avoid the detrimental effects
of abiotic stress on plant growth and development.
This comparison of gas exchange characteristics revealed the
importance of these parameters to predict fig performances in a
given environment. This work highlights the need to carry out other
studies in different environments and years in order to understand
better physiological fig tree behaviour under a range of conditions.
Declaration of Competing Interest
The authors declare that they have no conflict of interest.
Acknowledgements
We gratefully acknowledge the experiment station staff and
‘Water and Soil lab’ staff of the Regional Research Centre on Horticul-
ture and Organic Agriculture (CRRHAB) for their help to conduct
these experiments.
 A. AMMAR et al. / South African Journal of Botany 132 (2020) 30
37
References
Andersen, P.C., 1989. Leaf gas exchange characteristics of eleven species of fruit
crops in north Florida. Proceedings of the Florida State Horticultural Society
102, 229–334.
Angelopoulos, K., Dichio, B., Xiloyannis, C., 1996. Inhibition of photosynthesis in olive
trees (Olea europea L.) during water stress and rewatering. Journal of Experimental
Botany 47 (8), 1093–1100.
Bewick, V., Cheek, L., Ball, J., 2004. Statistics review 9: one-way analysis of variance.
Critical Care 8 (2), 130–136.
Can, H.Z., Hepaksoy, S., Akoy, U., Kutlu, E., 2000. Leaf characteristics and net gas
exchange of fig cultigens adapted to different climatic zones. Acta Horticulturae
516, 131–137.
Can, H.Z., Aksoy, U., 2007. Seasonal and diurnal photosynthetic behavior of fig (Ficus
carica L.) under semi-arid climatic conditions. Acta Agriculturae Scandinavica,
Section B - Soil & Plant Science 57 (4), 297–306.
Can, H.Z., Meyyaci, K.B., Balci, B., 2008. Determination of gas exchange capacity of some
Breba fig cultivars. Acta Horticulturae 798, 117 112.
Clifford, S.C., Kadzere, I., Jones, H.G., Jackson, J.E., 1997. Field comparisons of photosyn-
thesis and leaf conductance in Ziziphus mauritiana and other fruit tree species in
Zimbabwe. Trees Structure and Function 11, 449–454.
Du, Y., Zhao, Q., Li, S., Yao, X., Xie, F., Zhao, M., 2019. Shoot/root interactions affect soy-
bean photosynthetic traits and yield formation: a case study of grafting with
record-yield cultivars. Frontiers in Plant Science 10, 1–14.
Giorgi, F., Lionello, P., 2008. Climate change projections for the Mediterranean region.
Global and Planetary Change 63 (2-3), 90–104.
Gonz
alez-Rodríguez, A.M., Peters, J., 2009. Strategies of leaf expansion in Ficus carica
under semiarid conditions. Plant Biology 12, 469–474.
Greer, D.H., Weedon, M.M., 2012. Modelling photosynthetic responses to temperature
of grapevine (Vitis vinifera cv. Semillon) leaves on vines grown in a hot climate.
Plant Cell & Environment 35 (6), 1050–1064.
Greer, D.H., 2014. Seasonal changes in the photosynthetic response to CO2 and temper-
ature in apple (Malus domestica cv. ‘Red Gala’) leaves during a growing season with
a high temperature event. Functional Plant Biology 42, 309–324.
€rtensteiner, S., 2014. Different
Guyer, L., Hofstetter, S.S., Christ, B., Lira, B.S., Rossi, M., Ho
mechanisms are responsible for chlorophyll dephytylation during fruit ripening
and leaf senescence in tomato. Plant Physiology 166 (1), 44–56.
37
Hailemichael, G., Catalina, A., Gonzalez, M.R, Martin, P., 2016. Relationships between
water status, leaf chlorophyll content and photosynthetic performance in Tempra-
nillo vineyards. South African Journal for Enology and Viticulture 37 (2), 149–156.
Higgins, S.S., Larsen, F.E., Bendel, R.B., Radamaker, G.K., Bassman, J.H., Bidlake, W.R.,
Al Wir, A., 1992. Comparative gas exchange characteristics of potted, glasshouse-
grown almond, apple, fig, grape, olive, peach and Asian pear. Scientia Horticulturae
52 (4), 313–329.
Kutlu, E., Can, H.Z., Aksoy, U., Hepaksoy, S., 2000. Evaluation of gas exchange capacity
and physiological responses of selected Sarilop (Caliyrma) fig clones. Acta Horticul-
turae 517, 59–63.

Mlinari
c, S., Antunovi
c Duni
c, J., Stolfa, I., Cesar, V., Lepedus, V., 2016. High irradiation
and increased temperature induce different strategies for competent photosynthe-
sis in young and mature leaves. South African Journal of Botany 103, 25–31.
Palliotti, A., Silvestroni, O., Petoumenou, D., 2009. Photosynthetic and photoinhibition
behavior of two field grown grapevine cultivars under multiple summer stresses.
American Journal of Enology and Viticulture 60, 189–198.
Pige, L.C., Salager, J-L., Hossaert-Mckey, M., Roy, J., 2001. Carbon allocation to volatiles
and other reproductive components in male Ficus carica (Moraceae). American
Journal of Botany 88, 2214–2220.
Proietti, P., Famiani, F., 2002. Diurnal and seasonal changes in photosynthetic char-
acteristics in different olive (Olea europea L.) cultivars. Photosynthetica 40 (2),
171–176.
Psimisi, E., Vemmos, S.N., Mili, E., 2012. The photosynthetic activity and evaluation
of fruit quality in seven fig cultivars (Ficus carica L.). Acta Horticulturae 940,
341–348.
Urban, J., Ingwers, M.W., McGuire, M.A., Teskey, R.O., 2017. Increase in leaf temperature
opens stomata and decouples net photosynthesis from stomatal conductance in
Pinus taeda and Populus deltoides x nigra. Journal of experimental botany 68 (7),
1757–1767.
Vemmos, S.N., Petri, E., Stournaras, V., 2013. Seasonal changes in photosynthetic activ-
ity and carbohydrate content in leaves and fruit of three fig cultivars (Ficus carica
L.). Scientia Horticulturae 160, 198–207.
Vitale, L., Arena, C., Virzo De Santo, A., 2012. Seasonal changes in photosynthetic activ-
ity and photochemical efficiency of the Mediterranean shrub Phillyrea angustifolia
L. Plant Biosystems 146 (2), 443–450.
Zhang, S., Huang, W., Zhang, J., Cao, K., 2016. Differential responses of photosystems I
and II to seasonal drought in two Ficus species. Acta Oecologica 73, 53–60.