Agricultural Water Management 213 (2019) 749–759

Contents lists available at ScienceDirect

Agricultural Water Management

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

Impact of drought and salinity on olive water status and physiological T

performance in an arid climate
Lina Trabelsia,b, Kamel Gargouria, , Ameni Ben Hassenaa, Chaker Mbadraa, Mohamed Ghraba,
⁎

Bhekumthetho Ncubec, Johannes Van Stadenc, Radhia Gargourib

a
Olive Institute, University of Sfax, PO Box 1087, 3000, Sfax, Tunisia
b
National Engineering School of Sfax, University of Sfax, Route de la Soukra km, 4 - 3038 Sfax, Tunisia
c
Research Centre for Plant Growth and Development, School of Life Sciences, University of KwaZulu-Natal Pietermaritzburg, Private Bag X01, Scottsville, 3209, South
Africa

ARTICLE INFO ABSTRACT

Keywords: Effects of drought and salinity on water status, growth and physiological activity of olive can be temporary or
Olea europaea permanent and may impact olive production sustainability, especially in southern Mediterranean areas. Tunisia has a
Drought Mediterranean climate with high temperatures and low summer rainfall. Thus water stress problems are likely to be
Rainy period more severe for cultivated olive trees. In addition, the reduction in the availability of good quality irrigation water
Photosynthesis
will increase the use of saline water. Olive trees are able to tolerate low soil water availability and quality and
Olive water status
develop physiological adaptations to cope with water and salt stress. However, these adaptation strategies are limited
Olive recovery capacity
and permanent damages can be observed. The permanent effects caused to olive leaves due to drought, and the
capacity of irrigation with saline water to avoid these impacts is not well known. The aim of this work was to
compare olive leaves performance after a severe drought with, and without, irrigation and to assess recovering
capacities after a rainy period. Moreover, irrigation water quality effects were also evaluated. The results showed
that, photosynthetic rate was very low for rainfed plants, during drought, as compared to irrigated ones. After re-
watering, rainfed trees photosynthetic rate was only 55% of that of trees irrigated with fresh water. Irrigation with
saline water (EC = 7.5 dS m−1) reduced drought impact by increasing photosynthesis by 55% but remained lower
than that of fresh water by 23%. Thus olive leaves were unable to recover their whole photosynthetic capacity after
being exposed to severe water or salt stress. Furthermore, young leaves had the same photosynthetic capacity at the
beginning. This indicated that olive leaves lost permanently half of their photosynthetic activity during to drought
without irrigation. The use of saline water reduced this gap to 23% as compared to fresh water.

1. Introduction water deficit (Moriana et al., 2003, 2012; Ghrab et al., 2005, 2013;
Fernandez, 2013). Furthermore, drought involves alterations of mor-
The Mediterranean basin is the largest area in the world having phological and anatomical characteristics at the leaf level. It has been
specific climatic conditions suitable for olive cultivation. However, this reported that water stress frequently decreased leaf water status inducing
region is likely to face severe impacts due to climate change low leaf water potential and relative water content. This effect depends
(Tanasijevica et al., 2014). The Mediterranean region is expected to mainly on plant species and water stress severity (Aref et al., 2013). The
have, an overall reduction of annual precipitation of 39.1 ± 55.1 mm olive tolerates severe drought conditions with a series of adaptation
and an increase of air temperature of 1.57 ± 0.27 °C (from 0.84 to mechanisms (Chartzoulakis et al., 1999). These adjustments lead to
2.31 °C) inducing an increase of annual reference evapotranspiration of changes in leaf water status and stomatal closure, resulting in a reduction
92.3 ± 42.1 mm (Gargouri et al., 2010; Saadi et al., 2015). The agri- in photosynthetic rate. Hence, the leaf water status affects the efficacy of
cultural production has to adapt to this situation for sustainability (Ghrab photosystem II (PSII) activity and consequently the limitation of photo-
et al., 2008). Water shortage is known to induce many physiological synthesis (Fini et al., 2013). Loreto et al. (2003) and Centritto et al.
changes in plants (Aref et al., 2013). Indeed, the phenological stages of (2003) perceived that the reduction of photosynthesis could potentially
vegetative growth, flowering and fruit production are highly sensitive to be due to a decrease in both stomatal and mesophyll conductance.

⁎
Corresponding author.
E-mail address: [email protected] (K. Gargouri).

https://doi.org/10.1016/j.agwat.2018.11.025
Received 21 June 2018; Received in revised form 8 November 2018; Accepted 24 November 2018
Available online 02 December 2018
0378-3774/ © 2018 Elsevier B.V. All rights reserved.
L. Trabelsi et al. Agricultural Water Management 213 (2019) 749–759

Stomatal control is the major physiological factor in the optimization of technique (PRD) induced a slight reduction in pre-dawn and stem water
water use in drought conditions (Giorio et al., 1999; Yordanov et al., potential with the most important yield reduction was observed for the
2003; Lefi et al., 2004; Zlatev and Yordanov, 2004; Ben Hamed et al., rainfed (DRY) treatment. Oil content and quality improve with the in-
2016). Fernandez et al., (1997) and Moriana et al., (2002) noticed that creasing deficits for irrigated treatments, which resulted in the saving of
olive trees can reduce excessive water loss by closing their stomata. up to 50% of water requirements (Ghrab et al., 2005, 2013). Ghrab et al.
Consequently, the storage and diffusion of CO2 into the leaf is limited (2013, 2014) showed that PRD irrigation technique application could be
(Flexas et al., 2006). Biochemically, reduction of functional Rubisco or to used with low quality of water in sandy deep soil to improve productivity
inhibition of functional activity of PSII may limit photosynthesis (Lawlor without important accumulation of salts. Soil texture and Mediterranean
and Cornic, 2002). Changes in leaf anatomical characteristics are known climatic conditions affected the impact of saline water use management
to alter the CO2 conductance diffusion components from the substomatal for irrigation. The restriction by half (50%) of the water supply switched
cavities to the sites of carboxylation and thus contribute to the main- every month between wet and dry root zone caused a slight reduction of
tenance of photosynthetic rates even with the low stomatal conductance tree water status and cumulative yield (11%) over the four-year period of
(Chartzoulakis et al., 1999; Evans and Loreto, 2000). Thus, climate monitoring. Soil salinity presented seasonal variations affected by the ir-
change will result in an increase of irrigation demand (Masmoudi et al., rigation water applied. In this sandy soil salts were leached during the wet
2010). In these areas, agriculture faces enhanced land and water re- season and salinity of soil was kept to levels that were not harmful to olive
sources degradation; over-extraction of groundwater, soil salinization trees (Ghrab et al., 2013, 2014). Nevertheless, the efficacy of irrigation
and erosion (Ghrab et al., 2008). This is mainly due to the development with saline water to alleviate drought impacts on olive as compared to
of intensive orchard using poor water quality (Abdel Latef and Chaoxing, fresh water and rainfed conditions was not assessed.
2014). Indeed, olive is considered a moderately saline tolerant species. The actual work was conducted in the same orchard using rainfed
Due to scarce fresh water resources in the Mediterranean basin, irrigation and full irrigated (100% ETc) with saline water treatment (EC = 7.5 dS
with saline water (EC = 5–10 dS m−1) is often practiced. Nevertheless, m−1) in addition to a new treatment that corresponded to full irrigated
early rainfalls allow the removal of accumulated salinity from the root with fresh water (tap water EC = 2.46 dS m−1). It focused on the
zone which allows plant growth (Ghrab et al., 2013). Salinity affects possible effects of drought and water scarcity on olive tree water status
shoot growth and number of flowers and fruits, while the effects on yield and physiological activity. These effects can be temporary or perma-
depend on the salt concentration (Ghrab et al., 2013). Olive trees use nent, and may impact olive production sustainability especially in
several mechanisms of tolerance to salt accumulation. Salt tolerance is southern Mediterranean area. Olive trees are able to develop physio-
related to the capacity of olive to accumulate salt in the leaf vacuoles or logical adaptations to water stress. Although, these adaptation strate-
to the control of net salt import to the shoot (Chartzoulakis et al., 2002; gies are limited and permanent damages can be observed. Irrigation can
Ghrab et al., 2013). Salt stressed olive plants faces a reduction in me- overcome this shortage. Though, the permanent damages caused to
sophyll conductance to CO2 diffusion. This phenomenon is as rapid and olive leaves due to a severe drought and the capacity of irrigation with
reversible as for stomatal conductance. Moreover, the decrease of different water qualities to avoid these damages is not well known.
chloroplast CO2 content due to low stomatal and mesophyll conductance In this context, this study addresses the issues and impact of water
reduces photosynthesis, a process called diffusional limitations. shortage on olive trees by investigating the physiological behavior,
Under water stress, the potential to recover are a crucial component production and growth parameters of olive tree during and after
of a plant’s response (Miyashita et al., 2005). Indeed, the recovery ca- drought. The capacity of olive trees to recover their performance was
pacity determines plant establishment in the future. Many researchers also evaluated. Thus, the major aim of this work was to test how
have asserted that after re-watering olive recovers normal water status drought and salinity affected olive tree water status (RWC content, EL)
with leaf water potential and stomatal conductance similar to those of and leaf water potential estimation (Ψl), and the evolution of gas ex-
irrigated trees (Fernandez et al., 1997; Trentacoste et al., 2018). Con- change parameters such as Gs, transpiration (E) and photosynthetic rate
versely, stomatal conductance exhibited the slowest recovery. While the (Pn) in addition to leaf chlorophyll pigment content. Moreover, it fo-
mechanism is not well known, it has been linked to lower leaf water cused on the assessment of the impacts of severe drought in field con-
potential (Brodribb and Cochard, 2009) and to the accumulation of ditions on olive tree and the efficacy of irrigation with saline water
abscisic acid (ABA) during water withholding, which stands against (EC = 7.5 dSm−1) to mitigate it. Finally the recovery capacity of pho-
stomatal conductance recovery (Lovisolo et al., 2008). Photosynthesis tosynthetic activity after rainy period (re-watering by 68.8 mm of rain)
recovery depends on the rate and degree of photosynthesis inhibition was evaluated for rainfed and saline irrigation as compared to irrigation
during water shortage and also on plant species involved (Flexas et al., with saline water. For this reason three treatments were installed both
2009). It varies from rapid and complete recovery under moderate of them irrigated with tap (EC = 2.46 dS m−1) or saline water
stress to depressed or never complete after severe stress (Chaves et al., (EC = 7.5 dS m−1) and the third without irrigation. The drought se-
2009). Although olive tree response to water and salt stresses was verity and length were estimated using standardized precipitation
widely studied, the impacts of long drought, in field conditions, with index. The evaluation was based on leaf gas exchange, photosynthesis,
regards to temporary and permanent damages on leaf photosynthetic leaf water status (RWC and EL), pigments and shoots growth and pro-
activity quantification and capacity of irrigation with saline water duction. In addition, leaf water potential was estimated.
(EC = 7.5 dSm−1) to alleviate them were not assessed. These damages The objective of this work is to compare olive leave activities after a
are likely to reduce plant growth and production potential. severe drought with and without irrigation and to assess recovering ca-
Tunisia is among countries with limited water resources. An irrigation pacities after a rainy period (68.8 mm). In addition efficacy of saline
experiment was conducted in Sfax region (southern Tunisia) to evaluate water as compared to fresh water was assessed. Field experiment was
the combined effect of drought and irrigation with saline water on olive conducted on 25 years old olive trees (Olea europaea L. cv. Chemlali)
trees. Previous researches in the same experimental field than this work grown on sandy deep soil. This experimental design was installed in 2003
were carried since 2003 leaded to evaluate deficit irrigation approaches to assess efficacy of deficit irrigation on olive tree (Ghrab et al., 2013,
with the use of saline water (6.7 dS m−1), its impacts on olive tree and 2014) and was modified for the purpose of this work integrating fresh
adaptation mechanisms mainly physiological ones (Ghrab et al., 2005, water irrigation and drought assessment. The evolution of gas exchange
2013 and 2014). Three irrigation treatments (i) Control: full irrigated, (ii) parameters (Pn, E and Gs), WUE, chlorophyll (Chl), and plant water
DRY: rain-fed and (iii) PRD: partial root-zone drying were applied. In status (RWC) were followed and Leaf water potential (Ψl) was estimated
addition PRD was composed of two treatments: PRD30 and PRD15, which at the end of a severe drought period and after re-watering for rainfed
received 50% of ETc with an alternate irrigation switched every 30 and 15 and fully irrigated trees with tap and saline waters. Newly sprouted
days, respectively (Ghrab et al., 2005). Partial root-zone drying irrigation leaves activities were considered as initial situation for every treatment.

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2. Materials and methods overlapping the climatic demand and filling a part of water holding
capacity. Thus, since September 2016 re-watering was achieved by
2.1. Experimental site and water irrigation approach natural rainfall (68.8 mm). 10 trees were drip irrigated with tap water
(TW) while 10 trees were drip irrigated with saline water (EC = 7.5 dS/
The experiments were conducted in an orchard at the Olive Institute m) and the third treatment was composed by 10 trees under rain fed
in the region of Sfax (34°43’N, 10°41’E). Three irrigation treatments conditions and. Measures were taken after severe drought of 10 months
were installed in March 2015 and modified an experimental design in 16 July 2016, after re-watering in 24 October 2016 for adult and
installed in 2003 (Ghrab et al., 2013) by adding irrigation with fresh young leaves. The evolution of gas exchange parameters (Pn, E, and
water. There was no properly re-watering treatment was applied but Gs), WUE, chlorophyll (Chl), and plant water status (RWC) were fol-
rewatering was done by 68.8 mm rain that filled soil water holding lowed and Leaf water potential (Ψl) was estimated.
capacity of 5.5% (field capacity and wilting point of 8.8% and 3.3%
respectively) (Ghrab et al., 2013), The region is characterized by an arid 2.1.2. Plant sampling and experimental manipulation
climate of Mediterranean type, sandy deep soil (49.7% sand, 7% clay, Field experiment was conducted on 26 years old olive trees (Olea
43.3% silt). The mean annual precipitation and reference evapo- europaea L. cv. Chemlali) grown on sandy deep soil. 10 trees were drip
transpiration (ETo) were 220 and 1383 mm, respectively with marked irrigated with tap water TW (EC = 2.46 dS/m) while 10 others were
dry summers. The orchard was planted with twenty five-year-old olive under rain fed conditions (DRY) and also 10 trees were drip irrigated
trees (Olea europaea L.) cv. Chemlali at 4 m x 4 m spacing (625 trees with saline water FI (EC = 7.5 dS/m). Measures were taken after severe
ha−1) and drip irrigated. Standard agricultural practices in the region drought of 10 months in 16 July 2016 and one month after heavy rain
were applied. Irrigation was supplied by two dripper lines on each row of 68.8 mm in 24 October 2016 for adult and young leaves. Three re-
of olive trees located 0.5 m from the trunk using low quality ground- plications were taken per treatment at each sampling date. Leaves were
water (EC = 7.5 dS m−1). Saline water composition was 1130 mg/L taken all around the trees and five shoots for every tree were selected to
Na+, 1112.3 mg/L Cl-, 24 mg/L K+, 2.16 mg/L NH4+, and 40.51 mg/L be similar in potential yield and canopy and considered for the different
NO3- and pH of 7.35. The ground-water table depth was below 2.5 m. In measurements, surveys and quantifying the effect of drought and sali-
order to assess the impact of the irrigation with the saline ground- nity on the evolution of gas exchange parameters (Pn, E, Gs), WUE,
water, a treatment irrigated with tap water (EC = 2.46 dS m-1) was chlorophyll (Chl) content, and plant water status (RWC, EL). Leaf water
added. Tap water composition was Na+ 367.5 mg/L, Cl- 592.07 mg/L, potential (Ψl) was estimated using Gs. One month after rainfall mea-
K+ 13 mg/L, NH4+ 0.504 mg/L and NO3- 2.356 mg/L and pH of 8.02. surements were done on adult leaves and new leaves. These later served
to assess the leaf capacity before facing stresses.
2.1.1. Treatments applied and re-watering technique
The actual experiment design with three irrigation treatments was 2.1.3. ETc determination and water supply
installed in March 2015: Reference evapotranspiration (ETo) was calculated with Penman-
Monteith equation using daily data from a nearby automatic weather
1 Rain-fed: Olive trees were grown under rainfed conditions without station. The seasonal ETo values varied from 53 to 165.4 mm with a total
any irrigation supply. of 1382.7 mm (Table 1). The Crop evapotranspiration (ETc) was esti-
2 FI: Fully irrigated that applied daily the necessary saline water mated using FAO method applying the crop coefficient (Kc) for olive
(EC = 7.5 dSm−1) to match crop evapotranspiration ETc. trees FAO-56 (Allen et al., 1998). The monthly ETc varied from 48 to
3 TW: Fully irrigated that applied daily the necessary tap water 105 mm with frequent warm months. Effective precipitation (Ep), con-
(EC = 2.46 dSm−1) to match crop evapotranspiration ETc. sidered as 80% of total precipitation, ranged between 0 and 41.2 mm
monthly. Crop evapotranspiration (ETc) of the olive trees was calculated
The drought was evaluated (starting, end and severity) using SPI using the formula ETc = Kc * ETo, where ETo is the reference evapo-
(McKee et al., 1993, 1995; Soro et al., 2014). Basing on that length of transpiration calculated with the Penman-Monteith equation FAO-56
drought was determined. The end of drought was due to rainfall. During (Allen et al., 1998) and Kc is the crop factor with monthly values of 0.45
a severe drought, two treatments were irrigated to cover ETc. The first in July-August (drought period) and 0.65 on October-November (after
one was with saline water FI (EC = 7.5 dS m−1) and the second was rainy period). Daily weather data, recorded at the weather station of the
with tap water TW (EC = 2.46 dS m−1). A third treatment (rain-fed) National Institute of Meteorology (INM), located about 5 km from the
was installed without irrigation and did not receive other water than experimental site, were used to estimate ETo.
rain. The amount of received rainfall during a severe drought was very
low keeping the soil dry. Indeed, soil water holding capacity was 5.5%. 2.1.4. SPI determination
When considering a depth of 60 cm the required amount of water to fill The environmental conditions in the study area were characterized
this capacity is 429 m3/ha corresponding to 43 mm. Since October 2015 by a severe water deficit. Standardized Precipitation Index (SPI) was
the receiving rain water was less than the ETc leading to soil drying. calculated monthly from October 2015 to January 2017 to estimate
Thus during all the severe drought period water was not available in meteorological drought (McKee et al., 1993, 1995). SPI aims to describe
soil. At the end of drought (September 2016), 68.8 mm were received the major events of different types of drought observed in our study

Table 1
Monthly reference evapotranspiration ETo, culture coefficient Kc, monthly crop evapotranspiration ETc, effective precipitation Ep (as 80% of monthly precipitation)
and irrigation applied IA (mm) with irrigation treatments (FI : irrigated daily with 100% ETc with saline water (EC = 7.5 dSm−1); TW : irrigated daily with 100% ETc
with tap water (EC = 2.46 dSm−1) and rainfed condition) in the severe drought and after heavy rain period (68.8 mm) during experimentation from September 2015
to September 2016.
Sept. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sept. Total

ETo(mm) 113 87.3 61 53 59.2 71.2 98.3 113.2 137.8 160 165.4 148.4 114.7 1382.7
Kc 0.55 0.6 0.65 0.5 0.5 0.5 0.65 0.6 0.55 0.5 0.45 0.45 0.55 –
Etc (mm) 61.3 52.3 39.6 26.5 29.6 35.6 63.4 62 66 80 74.4 66.7 63 717.5
Ep (mm) 31 41.2 1.2 8.2 2.5 7.5 0 8.1 19 5 0 0 68.8 192.5
IA (mm) 30.3 11.1 38.4 0 0 28.1 63.4 53.9 47 75 74.4 66.7 0 533.7

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Table 2 2.4. Gas exchange and leaf water potential measurements

Categories of drought and wet defined by the values of SPI (McKee
et al., 1993). Gas exchange measurements were carried out on youngest and fully
SPI classes Degree of wet and drought expanded leaves, often healthy leaves and at the same physiological
age. The determination of leaf net CO2 assimilation rate (Pn) and sto-
SPI > 2 Extremely wet matal conductance (Gs) were determined on three leaves per plant with
1 < SPI < 2 Very wet
three replicate trees for each treatments from 9: 00 h to 13 h with a
0 < SPI < 1 Moderately wet
−1 < SPI < 0 Moderately dry LCA-4 portable photosynthesis system (ADC Bio Scientific Ltd., Hoddes-
−2 < SPI < -1 Severely dry don, UK). These measurements were made with the following specifi-
SPI < -2 Extremely dry cations: ambient CO2 concentration 390 μmol mol−1, average tem-
perature of leaf chamber was 39.2 °C. Average PAR at leaf surface was
1181.02 μmol m-2 s−1. Intrinsic water use efficiency (WUE) was cal-
area (Table 2). Precipitation occurred during autumn and winter sea- culated as the ratio of Pn over Gs (Maseyk et al., 2008). Leaf water
sons when the evaporative demand was low. Monthly applied irrigation potential (Ψl) was estimated and reported the average values and
water (IA) ranged between 11.1 and 75 mm for FI and TW treatments, standard deviation from relations with stomatal conductance (Gs) ac-
respectively, with annual total of 533.7 mm. The SPI, developed by cording to Wahbi et al. (2005); Guerfel et al., (2009), Boughalleb and
McKee et al. (1993, 1995) is only based on precipitation data. The index Hajlaoui (2010) and Masmoudi-Charfi et al. (2010):
has an advantage of being easily calculated, having modest data re- Ψl = (0.0087*Gs)-5.8545 (Boughalleb and Hajlaoui, 2010)
quirements, and being independent of the magnitude of mean pre- Ψl = (0.012*Gs)-4.8433 (Guerfel et al., 2009)
cipitation and hence it is comparable over a range of climatic zones. It is Ψl = (0.012*Gs)- 5.3079 (Wahbi et al., 2005)
calculated by fitting historical precipitation data. One of the significant Ψl = (0,001*Gs)-3,656 (Masmoudi-Charfi et al., 2010)
advantages of this index is that it can be calculated for different time-
scales to monitor meteorological droughts with respect to severity,
duration, onset, extent and end. Actual research based on the calcula- 2.5. Determination of chlorophyll and carotenoids concentration
tion of Standardized Precipitation Index (SPI) on a rainfall time series of
27 years from 1980 to 2017. The main input parameter is the monthly For total chlorophyll and carotenoid analyses, five leaf discs of fresh
rainfall in mm to provide a more representative month as possible of leaves of comparable physiological age (adult and young), were ground
the variable climatic conditions. The SPI index is expressed mathema- in 5 ml of 100% acetone solution, using a pestle and a mortar. After
tically as the ratio of difference between the annual rainfall of the year i filtration, extracts were adjusted to 20 ml with 100% acetone solution
(Pi) and the average rainfall of the series on the timescale considered and pigment contents were determined spectrophotometrically ac-
(Pm), and Standard deviation of the series on the timescale considered cording to the method of Hartmut and Buschmann, (1987).
(S) (Soro et al., 2014).

SPI = (Pi-Pm) / S 2.6. Leaf mineral content analyses (Na+, Cl−)

Samples for leaf mineral contents were collected in July 16th 2016.
Mineral content analyses were carried out on dry leaves obtained after a
2.2. Plants water status
drying at 70 °C until weight stabilization. The mineralization of these
samples was achieved by incineration at 450 °C and dissolution of ob-
Relative water content (RWC) was measured on youngest and fully
tained ash in 1 N nitric acid solution (HNO3−). Samples were processed
expanded leaves. Five leaves per plant of a similar age and position
using standard procedures (AOAC, 1990). The sodium (Na+) content
were detached with the petiole. Three replicate trees were used per
was measured using the flame emission photometry (Jenway PFP7,
each treatment at each sampling date during drought (16 July 2016)
Bibby Scientific limited, Staffordshire, UK). Chlorides (Cl−) content
and one month after a following rainy period (24 October 2016). Leaves
was determined by titration with 0.1 N silver nitrate (AgNO3) in the
were excised before dawn, weighed fresh (FW) and were immediately
presence of potassium dichromate according to a modified colorimetric
immersed in distilled water in a glass tube, and immediately sealed.
method of Mohr (Mathieu and Pieltain, 2003). All the values reported
After 48 h in dim light, the leaves were weighted to obtain the turgid
represent the means of at least three replicates and were expressed as
weight (TW). The dry weight (DW) was then measured after oven
percentage of dry weight (DW). For statistical analysis of data, each
drying at 75 °C for 48 h, and RWC was calculated as described by
three average value was considered as a replicate.
Laouar (1977):

RWC (%) = [(FW-DW)/ (TW-DW)] x 100

2.7. Shoot growth

Growth was measured at bud break and then monthly until the end
2.3. Electrolyte leakage (EL) of the growth season (pre-dormancy stage). Shoots were tagged at the
beginning of the study (January 2016) when trees were in dormancy
Electrolyte leakage (EL) values were measured as described by Lutts and shoot growth was measured from March to October 2016. Growth
et al. (1996). Fresh leaf samples were washed with distilled water to measurements of current-year shoots were made on eight healthy
remove surface-adhered electrolytes. Each leaf was cut into nine discs shoots from three trees of each treatment placed at different directions
of equal sizes and put into test tubes containing 10 ml distilled water around the canopy. The lengths of all apical and axillary shoots were
and incubated at room temperature on a rotary shaker for 24 h. Sub- measured.
sequently, the initial electrical conductivity of the medium (EC1) was
assessed. The samples were placed in an oven (90 °C) for 30 min. Then,
they were cooled at 25 °C and the second electrical conductivity (EC2) 2.8. Olive yield
was measured. Electrolyte leakage (EL) was calculated using the fol-
lowing formula: Yield per tree was measured at harvest on 10 trees per treatment.
Olive fruits were harvested by hand on December 2016. Olive yield was
EL (%) = (EC1/EC2) x 100 recorded and average production was reported in kg/tree.

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2.9. Statistical analysis

One way ANOVA analysis was applied to examine water availability

and quality effects on leaf water status shoot growth and gas exchange
using SPSS 13.0 for windows (SPSS Inc. Chicago, IL, USA). Significantly
different means were separated using the Duncan’s Multiple Range Test
at P < 0.05 significance level.

3. Results

The climate at the experimental site is basically an arid

Mediterranean type, characterized by: low minimal temperature at the
beginning of the crop cycle, on average 3 °C for January and February
2016; high maximum temperature of about 39 °C in summer during
olive growth, low and irregular rainfall, with an accumulated annual Fig. 1. Leaf relative water content (%) in olive trees cv. Chemlali Sfax under
value of around 123.7 mm, and a high evaporative demand, with an three treatments: Rain-fed conditions (Rain-fed), Full Irrigation with saline
accumulated annual evapotranspiration around 1382.7 mm. water (FI) and Full Irrigation with tap water (TW), evaluated during drought
and after rainy period for adult and young leaves. Data are means ± S.E. of 3
independent replicates. Different letters (a, b and c) indicate significant dif-
3.1. Drought occurrence
ferences at the 5% level between periods within the same treatment. Capital
letters (A, B and C): indicates significant differences at the 5% level between
SPI was used to determine length and severity of drought and cal- treatments within the same period.
culated in order to distinguish between dry and wet months and the
drought occurred when a succession of three dry months was registered
(Ben Abdelmalek et al., 2016). Thus, drought conditions are accepted 3.2.2. Electrolyte leakage (EL)
when the SPI was below -1 and ended when the SPI becomes positive. Electrolyte leakage is considered as a reliable indicator of the oxi-
The classes of SPI index are subdivided into moderate, severe and ex- dative stress resulting from abiotic constraints (Ben Hamed et al.,
treme for both dry and wet periods as shown in Table 2. 2012). The measurement of electrolyte leakage in leaves from plants
SPI indicated a long dry period from October 2015 to August 2016. subjected to different treatments showed that EL was significantly
Indeed, it ranged between -1 and 0 for nine months indicating moderate higher under rain-fed conditions during drought (76.96%) as compared
drought while the drought was severe for two months on 2015 such as to FI (62.75%) and TW (61.09%) with differences between irrigated
December (-1) and March (-1,06) (Table 3). After that dry period, trees at the same sampling date (Fig. 2). After heavy rainfall (68.8 mm)
September 2016 was classified as moderately wet with SPI of 0.63 the EL decreased for all the treatments. However, EL was significantly
followed by 68.8 mm of rain a moderately dry month and then one higher for rain-fed treatment and TW (39.74 and 37.22%) as compared
moderately wet (November) and one extremely wet (December 2016) to FI (29.82%) in mature leaves. Conversely, no significant differences
with SPI of 2.79. Based on these results the drought period was from were detected between treatments for leaves sprouted after rainy period
October 2015 to August 2016. For this reason two periods were iden- and EL ranged between 29.82 and 31.20% for all the treatments. On the
tified and were used for physiological records and sampling for bio- other hand, the decrease of EL was gradual for TW where highest EL
chemical analyses. These two periods were beginning of October 2015 value was recorded during drought followed by that recorded in mature
to August 2016 that corresponded to the last month of a long drought leaves after re-watering and the lowest was that of young leaves. In
and the end of October 2016 that corresponded to the end of drought contrary the EL drop was high for Rain-fed and FI. For these treatments
and one month after re-watering by 68.8 mm of rainfall in September the highest values were those of mature leaves during drought and the
2016. lowest values were those recorded after receiving 68.8 mm of rain
equally for young and mature leaves.
3.2. Effects of drought on plant water status
3.3. Effects of drought and re-watering on leaf water potential, gas
3.2.1. Relative water content (RWC) exchange and intrinsic water use efficiency
Since the leaf relative water content (RWC) allows determining the
actual leaf water retention under unlimited water supply, it can indicate 3.3.1. Leaf water potential (Ψl)
water status in plants (Li et al., 2010). The determination of the RWC in Leaf water potential remained almost constant for mature leaves for
leaves of olive plants subjected to the different water treatments every treatment before and after rain (Table 4). However, irrigated
showed significant differences (p ≤ 0.05) during drought between treatments had a converse trend for young leaves as compared to rain-
rainfed and irrigated trees independently of water quality FI and TW. fed trees. Indeed, young leaves showed lower Ψl than mature ones in
Rainfed trees had significantly lower RWC (66.64%) compared to FI irrigated conditions, while Ψl was higher for young leaves in rainfed
(83.08%) and TW (80.99%). However, after rainy period no significant conditions. During the severe drought period, Ψl of rain-fed trees was
differences were detected between treatments with values ranging be- -4.86 MPa, that was significantly lower than that of FI trees (-3.8 MPa),
tween 77.74 and 80.5% for young leaves and 78.27 and 82.25% for which was significantly lower than TW (-3.48 MPa). After re-watering
mature leaves (Fig. 1). Thus RWC was almost the same independently by 68.8 mm of rain, Ψl did not show any significant variation but it
from the type of leaves adult or young. decreased slightly for Rain-fed (-4.5 MPa) and FI (-3.66 MPa). Rain-fed

Table 3
SPI evolution from October 2015 to September 2017.
SPI Oct. Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sept.

2015 / 2016 −0.45 −0.85 −1 −0.65 −0.62 −1.06 −0.51 −0.22 −0.1 −0.32 −0.48 0.63
2016 / 2017 −0.39 0.28 2.79 −0.57 −0.07 −0.68 0.23 −0.63 0.05 −0.32 −0.48 −0.82

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3.3.2.2. Net photosynthesis rate (Pn). During drought, Pn was

significantly lower as compared to irrigated treatments (Table 4).
Indeed, Pn of rainfed treatment was 3.92 μmol CO2 m−²s-1 while it
was 15.47 and 19.41 μmol CO2 m−²s-1 for FI and TW respectively. After
receiving 68.8 mm of rain, Pn increased equally for mature and young
leaves in rainfed conditions reaching 10.09 and 12.15 μmol CO2 m−²s-1,
respectively. Conversely Pn of FI trees did not vary after re-watering
while it was higher in mature leaves (21.9 μmol CO2 m−²s-1) than
young leaves (12.46 μmol CO2 m−²s-1) for TW. Moreover, in irrigated
treatments photosynthetic rate was the same independently of water
quality and leaf age. Irrigation with saline water reduced the drought
impact by increasing Pn by 60% but remained under that with TW
treatment by 20% during severe drought. However, after rain, the
improvement of leaf water status was accompanied by 55% recovery
capacity of photosynthetic activity for rain-fed treatment, reaching
Fig. 2. Electrolyte leakage (%) in olive trees cv. Chemlali Sfax under three values of 10.09 μmol CO2 m-2s-1. While Pn in FI were significantly
treatments: Rain-fed conditions (Rain-fed), Full Irrigation with saline water (FI) (p ≤ 0.05) similar to the irrigated plants with tap water at the same
and Full Irrigation with tap water (TW), evaluated during drought, after rainy period. The relationship between photosynthetic rate (Pn) and stomatal
period for adult and young leaves. Different letters (a, b and c) indicate sig- conductance (Gs) (Fig. 3) indicates that all treatments maintained a
nificant differences at the 5% level between periods within the same treatment. high Pn and high Gs during severe drought and after re-watering
Capital letters (A, B and C): indicates significant differences at the 5% level periods.
between treatments within the same period.
3.3.2.3. Transpiration (E). During drought, transpiration was
trees had significantly lower Ψl than the other treatments. Young leaves significantly lower for rained treatment (0.61 mmol H2O m−2s-1)
had the same Ψl for all the treatments. It ranged between -4.02 (Rain- compared to irrigated trees. However, trees irrigated with saline
fed) and -4.3 MPa (TW). water had lower E (6.08 mmol H2O m−2s-1) than those irrigated with
tap water (8.47 mmol H2O m−2s-1). After heavy rain (68.8 mm), rainfed
mature leaves had significantly lower E than irrigated treatments while
3.3.2. Leaf gas exchange parameters no differences were detected for young leaves. Moreover, E decreased
3.3.2.1. Stomatal conductance (Gs). Stomatal conductance (Gs) during a after re-watering for irrigated treatments and the decrease was marked
drought period ranged between 8.24 mmol H20 m−2s-1 in rainfed plants for young leaves while it increased for rain-fed treatment.
and 170.34 mmol H20 m−2s-1 for TW respectively. Rain-fed treatment
was significantly lower than FI that was significantly lower than TW. 3.3.3. Intrinsic water use efficiency (WUE)
After heavy rain of 68.8 mm, young and mature leaves showed different The intrinsic water use efficiency, exhibited significant differences
behavior. In mature leaves, Rain-fed treatment had significantly lowest (p ≤ 0.05) between rained plants and irrigated ones (FI and TW) during
Gs (48.94 mmol H20 m−2s-1) as compared to the irrigated treatments, drought periods. During the water stress, rainfed trees maintained the
which had Gs ranging between 147.7 (FI) and 158.1 (TW) mmol highest WUE 0.56 μmol CO2 / mmol H2O. During severe drought WUE
H20 m−2s-1. Conversely, no significant differences were observed of FI and TW were 90 and 80% respectively higher than Rain-fed
between treatments in young leaves for Gs that varied between treatment. However, after heavy rain of 68.8 mm no significant differ-
72.92 mmol H20 m−2s-1 (TW) and 105.91 mmol H20 m−2s-1 (rain-fed). ences (p ≤ 0.05) were detected between treatments. During this period
Gs did not show significant differences between drought period and after WUE ranged between 0.12 and 0.26 μmol CO2 / mmol H2O.
receiving rain (68.8 mm) in mature leaves for all the treatments.
However young leaves had higher Gs for rainfed treatment as 3.3.4. Total chlorophyll Chl (a + b) and carotenoid contents(Car)
compared to mature leaves before and after re-watering. Conversely Gs Significant effects of water status and irrigation method were no-
of young leaves was lower than mature leaves for irrigated treatments. ticed on leaf total Chlorophyll (Chl a + b) contents. Indeed, during

Table 4
Leaf water potential (Ψl) (MPa), Net photosynthesis (Pn) (μmol of CO2 m−2 s-1), stomatal conductance (Gs) (mmol of H20 m−2 s-1), transpiration rate (E) (mmol of
H20 m−2 s-1) and water use efficiency (WUE) (μmol of CO2 m-² s-1 / mmol of H2O m−2 s-1) for mature and young leaves of the three treatments: Rain-fed conditions
(Rain-fed), Full Irrigation with saline water (FI) and Full Irrigation with tap water (TW), evaluated during drought and after rainy period.
Drought After rainy period

Mature leaves Young leaves

Rain-fed FI TW Rain-fed FI TW Rain-fed FI TW

Ψl −4.85 ± 0.06 −3.8 ± 0.13 −3.48 ± 0.15 −4.5 ± 0.15 −3.66 ± 0.22 −3.58 ± 0.22 −4.02 ± 0.49 −4.2 ± 0.07 −4.3 ± 0.41
aA bB bC aA bB bB bA aA aA
Pn 3.92 ± 2.3 15.47 ± 0.03 19.41 ± 2.56 10.09 ± 2.59 17.09 ± 3.3 21.9 ± 4.75 12.15 ± 5.16 12.33 ± 2.79 12.46 ± 2.8
aA aB bB bA aB bB bA aA aA
Gs 8.24 ± 6.74 132.26 ± 15 170.34 ± 18.23 48.94 ± 18.26 147.7 ± 26 158.81 ± 26 105.91 ± 58.73 84.6 ± 8 72.92 ± 48.76
aA bB bC aA bB bB bA aA aA
E 0.61 ± 0.44 6.08 ± 1.2 8.47 ± 1.31 2.82 ± 0.78 5.27 ± 0.78 6.44 ± 1.59 3.65 ± 0.94 3.24 ± 0.4 3.31 ± 1.97
aA bB bC bA bB abB bA aA aA
WUE 0.7 ± 0.66 0.12 ± 0.01 0.11 ± 0 0.25 ± 0.1 0.12 ± 0.01 0.14 ± 0 0.12 ± 0.01 0.14 ± 0.02 0.26 ± 0.17
bA aA aA aA aA aA aA aA aA

Data are means ± S.E. of 3 independent replicates. Different letters (a, b and c) indicate significant differences at the 5% level between periods within the same
treatment. Different capital letters (A, B and C): indicate significant differences at the 5% level between treatments within the same period.

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Fig. 5. Carotenoids (mg /g FW) in olive trees cv. Chemlali Sfax under three
treatments: Rain-fed conditions (Rain-fed), Full Irrigation with saline water (FI)
and Full Irrigation with tap water (TW), evaluated during drought and after
rainy period for adult and young leaves. Data are means ± S.E. of 3 in-
Fig. 3. Changes in photosynthetic rate (Pn) in relation to stomatal conductance
dependent replicates. Different letters (a, b and c) indicate significant differ-
(Gs) for all treatments: Rain-fed, FI and TW and all periods.
ences at the 5% level between periods within the same treatment. Capital letters
(A, B and C): indicates significant differences at the 5% level between treat-
severe drought period, all plants maintained low chlorophyll content ments within the same period.
which ranged between 0.91, 0.87 and 1.24 mg/g for rain-fed, FI and TW
treatments, respectively, with no significant differences (p ≤ 0.05) between all the treatments and without differences between adult and
(Fig. 4). However, after heavy rain of 68.8 mm, chlorophyll content young leaves (Fig. 6). Leaf chloride concentration showed the same
increased for all the treatments and ranged between 2.68, 4.29 and trend during and after drought for the three treatments. Indeed, the
3.98 mg/g. The rain-fed plants showed significantly lower increase 66% lowest values were recorded during drought and increased after rain. In
in Chl content compared to irrigated treatments FI and TW. No sig- addition, young leaves had higher concentrations than adult ones. The
nificant effect of salinity on Chl content was observed in FI as compared lowest values were recorded for TW followed by FI and rainfed treat-
to TW treatments. Conversely, the young leaves of the three treatments ments.
had the same chlorophyll content about 4.5 mg/g.
During the severe drought period, Rain-fed and FI trees showed the
3.3.6. Effect of drought and salinity on plant growth, development and
lowest leaf carotenoid contents ranging between 0.19 and 0.20 mg/g
production
(Fig. 5.). Conversely, TW had significantly (p ≤ 0.05) higher carotenoid
The elongation of olive shoots subjected to different water treat-
content reaching 0.29 mg/g. No changes were observed after re-wa-
ments are shown in Fig. 7. Rain-fed treatment led to very low shoot
tering in mature leaves for all the treatments. In contrary young leaves
growth (0.2 cm) and production (5.76 ± 12.4 kg/tree) during the
exhibited the same carotenoid contents for all the treatments of about
whole experimentation period as compared to FI and TW treatments.
0.29 mg/g.
Saline water irrigation (EC = 7.5 dS/m) slowed significantly (p ≤ 0.05)
growth rate during severe drought period as compared to tap water
3.3.5. Leaf mineral content (Na+, Cl−) treatment (TW). However, FI induced a significant growth increase as
Both sodium and chloride concentration increased for FI and in compared to rain-fed treatments accompanied with a higher production
Rain-fed conditions. Leaf Na concentration was higher during drought (16 ± 6.75 kg/tree). Thus, irrigation with saline water reduced the
for FI (0.7%) than tap water (0.21%) and Rain-fed (0.34%) (Fig. 6). drought impact by increasing elongation by 90% but remained lower
After rainy period the same trend was observed with the highest Na that with TW by 50%. TW had lower production (10 ± 8.6 kg/tree)
contents for rainfed followed by FI and TW with significant differences than FI. A gradual increase of growth was marked in plants irrigated
with saline water (FI) and tap water (TW) to reach by the end of ex-
perimentation 9.03 and 9.7 cm, respectively. After heavy rain
(68.8 mm), irrigated trees with saline water had more important growth
level similar to TW. Thus, by the end of the experimentation, both of
the irrigated treatments have the same shoot growth with no significant
differences (p ≤ 0.05).

4. Discussion

4.1. Effects of drought and saline water on olive gas exchanges

According to the results obtained in this study, the analyses of

photosynthetic parameters were affected by drought. Net photosynth-
esis (Pn) was highly altered in rainfed trees as compared to irrigated
Fig. 4. Chl (a + b) (mg /g FW) in olive trees cv. Chemlali Sfax under three ones. In fact, a significant (p ≤ 0.05) decrease was recorded in rainfed
treatments: Rain-fed conditions (Rain-fed), Full Irrigation with saline water (FI) plants about 3.92 μmol of CO2 m−2s-1. This coincided with a decrease
and Full Irrigation with tap water (TW), evaluated during drought and after in stomatal conductance (Gs) and the lowest values of leaf water po-
rainy period for adult and young leaves. Data are means ± S.E. of 3 in- tential (Ψl) during the drought period. The decrease in stomatal con-
dependent replicates. Different letters (a, b and c) indicate significant differ- ductance and photosynthetic rate due to water stress on is in agreement
ences at the 5% level between periods within the same treatment. Capital letters with other studies, which reported a similar reduction in leaf photo-
(A, B and C): indicates significant differences at the 5% level between treat- synthetic rate with the reduction in leaf stomatal conductance. A si-
ments within the same period. milar trend has also been reported in pistachio trees (Abbaspour et al.,

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morphological modification of intercellular space. This can also be due

to biochemical or cell membrane alterations. Therefore, the stomatal
limitation of photosynthesis, through stomatal closure, can be con-
sidered as a primary event followed by respective changes of the pho-
tosynthetic reaction as drought severity intensifies.
After re-watering water stress persistently affected photosynthetic
activity. Indeed, the increase of the assimilation rates of rain-fed trees,
after re-watering, indicated the improvement of intercellular CO2 con-
centration which was associated with a gradual recovery of photo-
synthetic capacity. However, this recovery was limited to 55% as
compared to FI and TW. Thus drought caused permanent damages to
leaf photosynthetic capacity. In addition, during drought and even after
rewatering FI mature leaves were not able be as efficient as TW ones.
However, young leaves had the same photosynthetic capacity in-
dependently from the treatment. This indicated that initially all the
leaves had the same capacity even for plant suffering drought or sali-
nity. Then drought or salt stress induced leaf capacity alteration and
even after re-watering they were not able to recover their initial per-
formance. Hence, drought and salt stresses alter the leaf capacity in an
irreversible way if their severity is very intense. It is important to notice
that the use of saline water mitigated the impact of drought. However,
this alleviation didn’t reached reach full capacity since a part corre-
sponding to 23% was lost due to salinity even after rewatering Without
irrigation olive leaves lost permanently half of their photosynthetic
activity during severe drought. The use of saline water (EC = 7.5 dS/m)
reduced this gap to 23% as compared to fresh water. This can be ex-
plained by many basic physiological and biochemical mechanisms at-
tributed to photosynthetic inhibition by water and salt stress intensity.
Indeed, during drought for rainfed and FI stomatal conductance was
reduced significantly reducing gas exchange and photosynthetic ac-
tivity. Diffusional resistances in the gaseous phase of CO2 transport
pathway from ambient air to carboxylation sites due to Gs and meso-
phyll damage were reported by Flexas et al. (2012) and Fernandez
(2013) in olive tree and Ben Hamed et al. (2016) in pistachio. After re-
watering raindfed trees were not able to fully recover Gs and conse-
Fig. 6. Na+ and Cl− contents in olive leaves for three treatments: Rain-fed, Full quently Pn. In contrary, FI trees recovered stomatal conductance while
Irrigation with saline water (FI) and Full Irrigation with tap water (TW) in July photosynthetic activity was not fully recovered as compared to FW.
2016. Data are means ± S.E. of 3 independent replicates. Different letters (a, b This can be explained by the fact that some impairment either of
and c) indicate significant differences at the 5% level between treatments.
photochemical or biochemical processes other than Rubisco (ribulose-
1,5-bisphosphate carboxylase/oxygenase) which greatly depends on
photoinhibition in olive. These have already been suggested by
Fernandez (2013) in olive tree and Sapeta et al. (2013) and Ben Hamed
et al. (2016) in pistachio. The impact of drought on stomatal con-
ductance indicates the failure of stomata to recover upon a rewatering
period. This suggests that the control mechanism of stomatal opening
and closing was affected by the water deficit imposed. Photosynthetic
recovery depends largely on previous water stress severity. The reg-
ulation of Gs after rewatering may be explained by a recovery of leaf
specific hydraulic conductivity.
The sequential decreases of Pn and PSII under drought are thought
to be caused by the close linkage between the activity of PSII and
photosynthesis (Chaves and Oliveira, 2004). This was most marked
when CO2 assimilation was limited by the decrease in stomatal con-
Fig. 7. Effect of drought and salinity on shoot growth and development of olive ductance Gs that can be considered as an important protective me-
trees cv. Chemlali Sfax for three treatments: Rain-fed, Full Irrigation with saline chanism in olive trees under drought and salt stress. This result was
water (FI) and Full Irrigation with tap water (TW), evaluated during drought
confirmed by a significant relationship between Pn and Gs. There was a
and after rainy period. Data are means ± S.E. of 24 measurements. Different
strong correlation; R² = 0.63, 0.74 and 0.76 for Rain-fed, FI and TW
letters (a, b and c) indicate significant differences at the 5% level between
treatments within the same period.
respectively.

4.2. Effects of drought and saline water on olive water status

2012; Ben Hamed et al., 2016), and olive trees (cv. Chemlali) (Guerfel
et al., 2009). This indicates that stomatal closure is one of the factors Intrinsic water use efficiency changed significantly with increased
limiting CO2 uptake under water shortage which was also found by drought. Indeed, constant Pn is estimated for decreasing Gs. This is a
Fernandez (2013). In fact, the reduction in CO2 assimilation in rain-fed key trait for the adaptation of olive to drought, and explains the higher
plants may reduce the leaf mesophyll conductance. Mesophyll con- WUE recorded in rainfed plants than irrigated ones. Irrigated plants had
duction is likely to be reduced due to tissues shrinkage causing constant stomatal conductance before and after rain converse to rainfed

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trees that had low Gs during drought and increased immediately after watering leads to a total resumption of initial water status which
re-watering but lower than that of irrigated trees. Therefore, irrigated showed that restoring plant water is reversible and easy to fully re-
trees showed better vegetative growth and yield. Thus even though cover. Moreover, olive tree water status was not affected by salinity
olive trees maintained high WUE under drought in rain-fed conditions stress but drought reduced leaf turgor even if all plants had a low water
physiological efficiency was very low, limiting plant growth and pro- potential. This indicated that the trees suffered water shortage and rain-
duction potential. fed plants were more efficient for water use for photosynthesis than
The leaf water status showed that leaf water potential (Ψl) de- irrigated ones. Therefore, the olive trees water status has been put in
creased significantly under severe drought for rainfed plants as com- evidence by RWC. The correlation between stomatal, mesophyll and
pared to irrigated ones for FI and TW treatments. Leaf water potential leaf hydraulic conductance, and the timing of each during regulation
was estimated using Gs and results were considered to draw trends and under severe drought, provided novel insights, showing that a decline
raise differences between treatments and not fully reported the water of leaf conductance is the earliest response to decreasing water avail-
status of leaves. The obtained results indicated that the status of irri- ability, and they propose that it triggers the later decline of stomatal Gs
gated trees was better than that of rainfed ones. In addition, it was and mesophyll conductance which reported also in rice by (Wang et al.,
lower for trees irrigated with saline water than those irrigated with 2018; Flexas et al., 2018) while the use of saline water (EC = 7.5 dS m-
fresh water. In fact, the minimum Ψl in the present study was -4.86 MPa 1) reduced drought impact by increasing water status (RWC) by 18%
registered for rainfed plants. This demonstrated the impact of dry and but remained similar to fresh water (EC = 2.46 dS m-1).
saline conditions on water status of olive tree. However, the observed
low leaf water potentials were not directly linked to leaf water status. 4.3. Effects on olive physiological performance
This decline revealed a similar reduction in leaf photosynthetic activity
which is in agreement with other studies (Perez-Martin et al., 2014). The analysis of studied physiological parameters may clarify the
Nevertheless, leaf RWC was low in rainfed trees as compared to different strategies of olive trees to respond to water and salt stress.
irrigated ones that maintained the same values. This may be the result Under water stress olive had moderate reduction in CO2 assimilation
of effective control of stomatal opening through chemical signals, rate, high WUE and decreased chlorophyll content with decreasing leaf
particularly the abscisic acid (ABA) synthesized by roots in response to water potential. Chlorophyll depletion can be attributed to the sensi-
the dehydration of soil moisture which was also considered by Kholová tivity of this pigment to increasing environmental stresses, especially to
et al., (2010). Further, maintaining RWC at 66.64% in rainfed trees, salinity and drought, which was also been reported by other researchers
under severe drought, should allow olive to maintain minimal cell (Moran et al., 1994; Younis et al., 2000; Guerfel et al., 2009). Water
turgidity, reflecting a great capacity of osmotic adjustment. Indeed, this shortage decreases the vegetative growth. The use of saline water re-
state will induce simultaneous gas exchange and Pn decline. Moreover, duced the impact of drought by 45% on shoot development but re-
Pn, Gs and RWC were significantly correlated. This illustrated the re- mained lower than that of plants irrigated with tap water (TW) after re-
sponse of Pn to decreasing RWC and Ψl induced by decreasing the water watering. Moreover, Irrigation with saline water (EC = 7.5 dS m−1)
supply to the roots. So, decreasing RWC decreased Gs and Pn, although reduced drought impact by increasing production by 64% but remained
at a small value of RWC, Gs may reach a minimum but Pn may continue higher than that of fresh water by 38%. Nevertheless, it is difficult to
to decrease (Lawlor and Cornic, 2002; Boussadia et al., 2008). draw conclusions for olive production since olive tree is alternate
RWC was relatively high for all the treatments during drought with bearing tree and at least two bearing seasons are necessary for that.
significantely lower values for rainfed treatment. After rewatering, all Negative effect of salinity was previously reported on pistachio (Ben
the treatments had the same values indicating for both cases a good cell Hamed and Lefi, 2015). Growth and development could be decreased
water status. This is suggesting modifications of leaf structure in re- with the increased degree of salinity and duration of imposed saline
sponse to water stress not only improve resistance of leaf cells to col- water irrigation when compared to tap water treatment (TW). Similar
lapse but also bring about changes in tissue elasticity. These adjust- findings were reported also on other species (Carr, 2014). Irrigation
ments have also been reported by Oertli et al., (1990) and Guerfel et al. with saline water (7.5 dS m−1 EC) reduced the ability of plants to take
(2009). Bacelar et al. (2006) found an increase in the cell wall elasticity up water as indicated by a significant limitation of transpiration and
in olive tree under low water conditions, which may reflect changes in low leaf water status, and this quickly caused reductions in growth rate
cell wall composition (Munoz et al., 1993). (Joly and Zaerr, 1987) ex- (Munns, 2002).
plained that more elastic cell walls can shrink more easily when sub- An accumulation of Na+ ions was observed in the tissues of all ol-
jected to stress, helping to maintain a higher turgor pressure and pro- ives tree during a severe drought period. As salinity increased, Na+
tecting cell walls from rupturing. This was confirmed by EL of 76.96, concentration in the leaves of all tested cultivar increased gradually.
62.75 and 61.08% for rain-fed, FI and TW respectively under severe Indeed, tissue sodium concentration was greater in leaves of trees ir-
drought. After rewatering, EL was reduced to a range between 29.82 rigated with saline water (EC = 7.5 dS m−1). Rainfed trees were clas-
and 39.74 for all treatments. On the other hand, rigid cells may help to sified second with regards to the concentration of Na in the leaves. This
maintain lower water potential at a given volume than elastic cells may be explained by the salt ascending from near saline water table due
(Patakas et al., 2002). This can result in an increase in the gradient in to soil desiccation during drought. The low values observed for TW are
water potential between the soil and the plant, thereby promoting more explained by salt leaching by fresh water. Na concentration in leaves
effective water uptake from drying soils and/or accelerating recovery was 0.33% in rainfed trees that corresponds to the level observed by
after watering (Bowman and Roberts, 1985; Guerfel et al., 2009). Ad- Kchaou et al. (2010) without toxicity on chemlali olive tree. However,
ditionally, cell membrane stability has been widely used to express the Na concentration was 0.7% for FI trees. This value is linked with
stress tolerance, and higher membrane stability could be correlated toxicity effects (Kchaou et al., 2010). TW had low level of Na in leaves
with abiotic stress tolerance (Bajji et al., 2001). In fact, our findings (0.21%) indicating that there was no salinity stress. Chloride con-
showed a significant increase in (EL) on rain-fed plants at severe centration in leaves of plant of the three treatments ranged between
drought period compared to irrigated ones (FI and TW). This may be 0.067 and 0.83%. These concentrations are within the normal range of
due to the fact that water shortage increases EL, also confirmed by Bajji concentration of Cl in olive leaves (Bartolini et al., 1991). However, the
et al. (2001). The comparison of the leaf water status during drought rainfed treatment had lowers concentrations indicating a limitation of
and after rewatering between treatments indicated that olive had good Cl absorption due to lack of water. In addition irrigation with saline
water status (RWC), low (EL) and for both of irrigated treatments before water did not affect the concentration of Cl in olive leaves. This may be
rain. Relative water content was kept high, even at maximum stress. It due to the limitation of transport of this element to leaves (Kchaou
increased after rain reaching those of irrigated ones. Therefore, the re- et al., 2010). The toxicity effects of Na on olive leaves may explain the

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L. Trabelsi et al. Agricultural Water Management 213 (2019) 749–759

reduction of gas exchange observed in FI treatment (Fernandez, 2013). Drainage Paper No. 56.
Olive shoot growth reduction under salt stress can be explained by Aref, I., El Atta, H., El Obeid, M., Ahmed, A., Khan, P., Iqbal, M., 2013. Effect of water
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