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Screening for PEG-induced drought stress tolerance in seedlings of durum

wheat (Triticum durum Desf.) cultivars

Article in Pakistan Journal of Botany · March 2021

DOI: 10.30848/PJB2021-3(5)

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Pak. J. Bot., 53(3): 823-832, 2021. DOI: http://dx.doi.org/10.30848/PJB2021-3(5)

SCREENING OF SEEDLINGS OF DURUM WHEAT (TRITICUM DURUM DESF.)

CULTIVARS FOR TOLERANCE TO PEG-INDUCED DROUGHT STRESS
AFEF OTHMANI*1§, SOUROUR AYED2§, ZOUBEIR CHAMEKH2, OLFA SLAMA-AYED3,
JAIME A. TEIXEIRA DA SILVA4, MOUNIR REZGUI5, HAJER SLIM-AMARA3 AND MONGI BEN YOUNES1
1
University of Carthage, Field Crops Laboratory, Regional Research Development Office of Agriculture in Semi Arid
North West of Kef, Tunisia
2
University of Carthage, Field Crops Laboratory, National Agricultural Research Institute of Tunisia,
Rue Hédi Karray 2049 Ariana, Tunisia
3
University of Carthage, Genetic and Plant Breeding Laboratory, Department of Agronomy and Biotechnology,
National AgronomicInstitute of Tunisia, 43, Avenue Charles Nicole, 1082 Tunis, Tunisia
4
Independent researcher, P. O. Box 7, Miki-cho post office, Ikenobe 3011-2, Kagawa-ken, 761-0799, Japan
5
University of Carthage, Science and Agronomic Techniques Laboratory , National Agricultural Research Institute of
Tunisia, Rue Hédi Karray 2049 Ariana, Tunisia
*Corresponding author’s email: othmaniafef@yahoo.com; §Equal conributors

Abstract

The effect of drought stress on 11 durum wheat (Triticum durum Desf.) cultivars was determined at the germination
stage. Cultivars were screened for drought tolerance. Six levels of osmotic stress (0, -0.47, -1.48, -3.02, -5.11 and -7.73 bars)
were assessed by applying different concentrations of polyethylene glycol (PEG-8000). There were significant differences
between treatments for all seedling characteristics (p<0.05, p<0.001), except mean daily germination (MDG). All seedling
traits also differed significantly (p<0.001) among all cultivars. In general, osmotic stress decreased seed germination
percentage, germination rate (GR), coleoptile length (CL), shoot length (SL), root length (RL), root/shoot (R/S) length ratio,
and root number (RN). Averaged over all osmotic stress levels, Mahmoudi had high MDG (0.55), GR (1.88), CL (4.20 cm),
SL (10.45 cm), and RL (9.93 cm), suggesting that this variety was highly tolerant to drought stress. There were high
correlation coefficients between different characteristics: SL had a positive and significant (p<0.01) correlation with CL (r =
0.83), RL (r = 0.74), and R/S length ratio (r = 0.67). This study showed that, based on morphological traits, preliminary
screening at an early stage for drought stress using PEG-8000 may facilitate the choice of an adequate cultivar for growth
under water stressed conditions.

Key words: Drought tolerance, Durum wheat, Germination, Osmotic stress.

Abbreviations: DGS, daily germination speed; GP, germination percentage; GR, germination rate; MDG, mean day
germination

Introduction caused by bioclimatic and farming conditions that are

more favorable in the north than in the center and the
In agricultural production, environmental stresses south of the country (Annabi et al., 2013).
play a significant limiting role (Ghader, 2014). Abiotic New challenges in modern agriculture include rapid
stresses present major challenges in sustaining crop yield population growth, climate change and the deterioration
(Chen et al., 2014). Salinity and drought are two common of arable lands, and enhanced agriculture is vital to face
environmental stresses that affect seed germination and global food demand with stress-tolerant crops present a
plant growth, especially in arid and semi-arid regions promising solution (Chen et al., 2014; Ulfat et al., 2017).
(Mohammadizad et al., 2013; Masondo et al., 2018). The development and release of new varieties that adapt
Drought stress decreases crop production while water to water deficit conditions could be a constructive way to
deficiency reduces plant growth and productivity surmount unsuitable environmental conditions. A good
(Castilhos et al., 2014; Gilani et al., 2020). Among understanding of factors limiting yield provides an
different stress factors, drought is ranked first, limiting opportunity to identify physiological traits that could
three quarter, or 454 million ha, of global crop production increase drought tolerance and yield under rainfed
(Kim et al., 2019). In Tunisia, where a gradient in severity conditions. Drought tolerance can be assessed in crops by
and frequency of drought exists from north to south, yield using physiological and agro-morphological tests, which
fluctuates considerably and is extremely low in dry years. serve as indirect selection criteria, thereby accelerating
Yield is also strongly associated with the amount of selection methods and hopefully resulting in cultivars
rainfall (Khakwani et al., 2012; Mansour & Hachicha, with increased yield and productivity under drought-
2014). In addition, most cereal cultivation lands are in the stressed climates (Ahmadizadeh, 2013).
northern to north-western areas where the climate varies The response of wheat cultivars to drought stress
from semiarid to dry sub-humid (Ferjaoui et al., 2014). has been examined extensively because soil drought
The national average yield of durum wheat is 14 ± 4 represents the main constraint for successful crop
qx/ha while that of bread wheat is 16 ± 5 qx/ha, but yield production. Plants can readily modify their metabolic
is higher in the north of Tunisia (18.4 ± 3.8 qx/ha) than at and physiological processes, as well as the morphology
the center and south (11.3 ± 4.7 qx/ha). This variation of the above-ground parts and the root system in
824 AFEF OTHMANI ET AL.

response to water deficit (Gallé et al., 2013; Castilhos In view of the importance of the germination phase
et al., 2014). Seed germination is the most sensitive and early plant formation under drought conditions, the
stage to abiotic stress (Jian et al., 2016) and involves present study was carried out to screen Tunisian durum
multiple morphological and physiological alterations wheat cultivars for drought tolerance by inducing water
take place during germination. In unfavorable stress by applying PEG at the germination and seedling
conditions, seeds may enter secondary dormancy to growth stages, and to quantify the association between
sustain their ability to germinate, and when conditions drought-associated seedling traits.
are favorable, such seeds are able to germinate. Under
semi-arid conditions, low humidity is a limiting factor Materials and Methods
during germination (Ahmadizadeh, 2013; Hafeez et al.,
2017; Li et al., 2018). Thus, there is a need to screen Plant materials: Grains of 11 durum wheat (Triticum
drought-tolerant genotypes in these areas (Mickky & durum Desf.) cultivars (Maâli, Mahmoudi, Om rabiaa,
Aldesuquy, 2017). Coleoptile length is an important Karim, Nasr, Salim, Maghrbi, Ben bechir, Souri,
factor controlling the emergence of seedlings from Agiliglabre, and Jnehkottifa) were obtained from the
deep sowing depths (Prabhakar et al., 2013; Lee et al., Genetic Laboratory of Cereal Crops, Agronomic Institute
2017). Seedling emergence is highly sensitive to water of Tunisia. Kernels were initially surface sterilized with
deficit and early drought hinders successful crop 12% sodium hypochlorite for 5 min and then rinsed twice
establishment by negatively affecting seedling with distilled water for 5 min each rinse. Ten grains of
emergence (Shahbazi et al., 2012; Hellal et al., 2018). each cultivar were placed in 90-mm diameter plastic Petri
Toklu et al., (2015) noted that seedling growth, dishes (Amazon, Orléans, France) on a single sheet of
coleoptile length and seed germination are filter paper (90-mm diameter; Fioroni, Paris, France).
fundamentals for successful stand establishment of Petri dishes (Sterilin Ltd., Cambridge, UK) were covered
yield plants. Studying the effects of water stress on the with lids to prevent the loss of moisture by evaporation.
growth and development of roots and shoots also has Grains were germinated under controlled conditions
merit as water stress is the most important abiotic in a dark growth room for 10 days, at 50% relative
constraint to increase grain yield in rainfed wheat- humidity and at an average day/night temperature of 22 ±
growing areas. Geneticists and breeders also breed 2°C (El Siddig et al., 2013). Seeds were considered to be
plants with root traits that improve productivity under germinated when the emerging radicle had protruded 5
drought (Comas et al., 2013; Shan et al., 2015). Wheat mm, at which point germinated seeds were removed
plants experience drought stress when the water supply (Prabhakar et al., 2013).
to roots becomes too little to support growth or when
transpiration becomes very high due to wind and Polyethylene glycol solutions: PEG (PEG-8000,
temperature (Bin Abdul Hamid, 2012). Germination molecular weight 8000 g mol-1, > 99.0% purity; Sigma-
and seedling characteristics such as germination
Aldrich, St. Louis, USA) aqueous solutions with different
percentage, germination rate, and seedling growth are
water potentials (-0.47, -1.48, -3.02, -5.11 and -7.73 bars)
the most viable criteria used for selecting drought
were established by dissolving 50, 100, 150, 200 and 250
tolerance in crops at the seedling stage (Jabbari et al.,
2013; Basha et al., 2015). The genotypic ability for a g of PEG in 1000 ml of distilled water. The control was
root/shoot length ratio contributes to drought tolerance, distilled water (0 bars). Water potential (Ψ) was calculated
which is genetically determined (Khan et al., 2020). by an equation that relates PEG concentration to water
The efficiency of soil water uptake by the root system potential (Michel, 1983):
is thus a key factor in determining the rate of
transpiration and drought tolerance (Ahmadizadeh, Ψ = 1.29 [PEG] [PEG]2T – 140 [PEG]2 – 4 [PEG]
2013; Tardieu et al., 2017). Roots capture water and
nutrients, besides anchoring the plant in the ground, where Ψ is the water potential of each treatment (bars),
and is logically seen as the most important organ to [PEG] is the concentration of PEG solution [g PEG (g
improve crop adaptation to water deficit (Vadez, 2014). H2O) -1] and T is temperature (°C).
The germination test in a high osmotic potential Five ml of each concentration of PEG solution, or
solution is one of the most important laboratory distilled water for the control, was added after every 2
methods to screen the drought tolerance of a crop plant days to a separate Petri dish to maintain the required
(Ahmadizadeh, 2013). At an early seedling stage, water potentials of each treatment constant.
polyethylene glycol (PEG) is used to induce plant
water deficit (Muscolo et al., 2014). PEG is a non-toxic Measurements: After 10 days of treatment, coleoptile
and non-penetrable osmotic agent that lowers water length (CL) was assessed by measuring the distance from
potential in mediumthat is employed to imitate drought the grain to the tip of the coleoptile in each replicate for
stress in plants (Toosi et al., 2014). PEG-6000 has all 10 seedlings (Prabhakar et al., 2013). Shoot length
often been used to induce water deficit and preserve (SL), root length (RL), ratio of root/shoot (R/S) length,
uniform water potential through an experimental period and root number (RN) were determined from five samples
(El Siddig et al., 2013). Seed germination as a function in each of the six humidity levels (five water potentials
of water potential is often tested with soil adjusted to plus one control) from each replication, based on
desired water potentials or by using PEG solutions established germination protocols (Moayedi et al., 2009;
(Prabhakar et al., 2013). Mohammadizad et al., 2013). In addition, the following
PEG-INDUCED DOUGHT STRESS TOLERANCE OF DURUM WHEAT 825

parameters were assessed daily: number of germinated stress levels (Table 1). These results corroborate those
seeds, germination percentage (GP), mean daily of Baloch et al., (2012), in which more than half of 16
germination (MDG; i.e., the mean number of seeds that spring wheat cultivars grown under osmotic stress did
germinated each day), daily germination speed (DGS), not show any reduction in seed germination even
and germination rate (GR). These parameters were though GP alone was not able to assess osmotic stress
calculated as follows: tolerance since it could not clearly discriminate the 16
cultivars.
GP = (total number of germinated grains / total number of
observed grains) × 100; Mean daily germination, daily germination speed and
MDG = final emergence/10 (Jajarmi, 2008); germination rate: There was a significant effect of water
DGS = 1/MDG (Maguire, 1962); potential on MDG and GR (Table 2). Prabhakar et al.,
GR = (n1× t1) + (n2 × t2) + (n3 × t3) + ... (ni × ti )/T
(2013) also observed that water potential influenced GR.
(Olmez et al., 2006):
Based on mean values, maximum GR and DGS were
where n = number of days in which grains germinated; t = observed in the control (0 bars). GR is significantly
number of germinated grains in each counting day; T = reduced when osmotic stress increases (Ezzat-Ahmadi et
total number of germinated grains. al., 2014). In contrast, osmotic stress caused an increase
in MDG. Abdoli & Saeidi (2012) also found that stress
Statistical analysis of data: A completely randomized caused by water deficiency augmented MDG of nine
design was performed in a factorial experiment with three wheat cultivars from 1.91 to 2.05 days. In 96 wheat
repetitions. The factors were durum wheat cultivars and genotypes, Dodig et al., (2014) showed that MDG
water potential. Analysis of variance (ANOVA) was increased after exposure to PEG-induced osmotic stress,
conducted for seed germination and growth measurements' delaying MDG by 14 days when osmotic potential was -
traits. The treatment means were compared by Duncan’s 0.4 MPa compared to the control treatment (distilled
multiple range test (p<0.05, p<0.01 and p<0.001) using water). Ahmad et al. (2017) claimed that such a response
SPSS software (version 16.0). is due to more time being required to germinate under
drought stress. There was also considerable variation in
Results and Discussion MDG, DGS and GR between cultivars (Table 2). Mujtaba
et al., (2016) found highly significant differences among
Emergence and related traits seed germination: The
six wheat genotypes for the same traits. Om rabiaa, which
germination percentage of 11 durum wheat cultivars as
a function of time under six water potentials is shown is considered to be tolerant to drought (Brini et al., 2007),
in Table 1 (on days 7, 8, 9 and 10, columns are vacant had the highest MDG and GR but a low DGS. Comparing
as no seeds germinated further on those days). Water the two new cultivars (Mâali and Salim), Salim had
potential and cultivar had a significant effect on seed significantly higher MDG and GR than Mâali (Table 3).
germination. Abido & Zsombik (2018) also noted that
Seedling growth
water potential affected the GP of seven Hungarian
wheat varieties. Shereen et al., (2019) found that GP Coleoptile length: The differences among water potential
was significantly reduced in eight rice genotypes as
treatments and among cultivars for CL were highly
water stress levels increased. Relative to the control,
significant, and there was a highly significant interaction
germination decreased as water potential with the
between water potential and cultivars (Table 4).
increase of water. Also, in this study, water stress
Prabhakar et al., (2013) reported that CL could affect seed
delayed and prevented seed germination. Water stress
at the germination stage could result in the delay, emergence since it was linked to emergence capacity from
decrease or even complete prevention of germination deep sowing depths, accounting for 60% of the
(Moral et al., 2015). differences between varieties in their study. High genetic
Differences in seed germination among cultivars variation was observed in CL. Numerous studies have
were observed. This genetic variation could be used to reported an association between long coleoptiles and
screen new wheat varieties adapted to unfavorable increased seedling emergence, weed restraint and
conditions in arid and semi-arid regions (Chachar et improved grain yield (Farhad et al., 2014). The shortest
al., 2014). Om rabiaa showed significantly higher seed CL was observed at the highest water potential (-7.73
germination (96.11%) than other cultivars under most bars). Mahmoudi had significantly longest coleoptiles
water potentials. Jnehkottifa had the lowest seed among all cultivars (Table 5). The tested cultivars
germination (42.78%). Except for Maâli, Souri, responded differently to osmotic stress (Table 5).
Agiliglabre and Jnehkottifa, all other cultivars showed Compared to normal conditions, CL was increased in
more than 80% seed germination at -5.11 bars. Maâli, Mahmoudi and Maghrbi at -0.47, -1.48 and -3.02
However, at -7.73 bars, only Om rabiaa and Nasr bars, in Karim, Salim, Agiliglabre and Ben Bechir at -
reached this range, indicating that the critical water 0.47 bars, in Nasr and Souri at -0.47 and -1.48 bars, and
potential for these cultivars lies between -5.11 and - in Jnehkottifa under high stress levels (-3.02 and -5.11
7.73 bars. Nevertheless, some cultivars did not show bars). Farhad et al., (2014) reported that water deficit
any decrease in seed germination over all osmotic could increase CL.
826 AFEF OTHMANI ET AL.

Table 1. Total seed germination (%) of 11 Tunisian durum wheat cultivars at different water potentials.

Water potential Days

Cultivar
(bars) 2 3 4 5 6 7 8 9 10
Mâali 53 67 73 80 83 nf nf nf nf
Mahmoudi 67 83 93 97 97 nf nf nf nf
Om rabiaa 73 100 100 100 100 nf nf nf nf
Karim 67 87 90 90 90 nf nf nf nf
Nasr 40 90 97 97 97 nf nf nf nf
0 Salim 33 97 97 97 97 nf nf nf nf
Maghrbi 63 87 90 93 93 nf nf nf nf
Ben Bechir 73 80 83 87 87 nf nf nf nf
Souri 47 67 70 73 73 nf nf nf nf
Agiliglabre 3 30 40 40 47 nf nf nf nf
JnehKottifa 0 37 43 43 53 nf nf nf nf

Mâali 73 73 77 80 80 nf nf nf nf

Mahmoudi 90 90 93 97 97 nf nf nf nf

Om rabiaa 93 97 100 100 100 nf nf nf nf

Karim 73 90 90 90 90 nf nf nf nf

Nasr 97 97 97 97 97 nf nf nf nf

-0.47 Salim 100 100 100 100 100 nf nf nf nf

Maghrbi 87 90 90 90 90 nf nf nf nf

Ben Bechir 47 63 63 63 63 nf nf nf nf

Souri 47 67 67 67 67 nf nf nf nf

Agiliglabre 7 57 60 63 63 nf nf nf nf

JnehKottifa 13 40 43 43 43 nf nf nf nf

Mâali 73 77 77 77 77 nf nf nf nf
Mahmoudi 83 83 87 93 93 nf nf nf nf
Om rabiaa 83 93 93 93 93 nf nf nf nf
Karim 90 100 100 100 100 nf nf nf nf
Nasr 67 97 97 97 97 nf nf nf nf
-1.48 Salim 67 97 100 100 100 nf nf nf nf
Maghrbi 73 77 83 83 83 nf nf nf nf
Ben Bechir 77 77 77 77 77 nf nf nf nf
Souri 33 47 50 50 50 nf nf nf nf
Agiliglabre 7 23 30 30 30 nf nf nf nf
JnehKottifa 7 30 40 40 40 nf nf nf nf
PEG-INDUCED DOUGHT STRESS TOLERANCE OF DURUM WHEAT 827

Table 1. (Cont’d.).

Water potential Days

Cultivar
(bars) 2 3 4 5 6 7 8 9 10
Mâali 57 67 73 73 73 nf nf nf nf
Mahmoudi 63 73 83 90 93 nf nf nf nf
Om rabiaa 97 100 100 100 100 nf nf nf nf
Karim 83 87 87 90 90 nf nf nf nf
Nasr 97 97 97 97 97 nf nf nf nf
-3.02 Salim 83 93 93 93 93 nf nf nf nf
Maghrbi 83 90 90 90 90 nf nf nf nf
Ben Bechir 67 70 70 70 70 nf nf nf nf
Souri 33 40 40 40 40 nf nf nf nf
Agiliglabre 13 30 37 43 43 nf nf nf nf
JnehKottifa 17 33 33 33 33 nf nf nf nf
Mâali 43 63 67 70 73 73 73 nf nf
Mahmoudi 73 87 97 100 100 100 100 nf nf
Om rabiaa 20 63 77 80 90 90 90 nf nf
Karim 40 67 87 87 93 93 97 nf nf
Nasr 40 67 87 87 93 93 97 nf nf
-5.11 Salim 10 67 80 90 90 93 93 nf nf
Maghrbi 0 47 60 80 90 93 93 nf nf
Ben Bechir 50 83 90 90 93 93 93 nf nf
Souri 47 63 63 63 63 63 63 nf nf
Agiliglabre 0 17 33 33 43 47 47 nf nf
JnehKottifa 0 27 30 40 40 43 43 nf nf
Mâali 10 27 37 40 50 53 53 nf nf
Mahmoudi 27 53 57 60 63 70 77 nf nf
Om rabiaa 0 33 67 70 90 90 93 nf nf
Karim 3 33 50 67 70 70 70 nf nf
Nasr 7 47 60 67 80 83 83 nf nf
-7.73 Salim 0 17 23 37 43 50 50 nf nf
Maghrbi 0 27 30 40 53 60 60 nf nf
Ben Bechir 13 37 40 43 50 53 53 nf nf
Souri 0 30 33 43 43 43 43 nf nf
Agiliglabre 0 13 33 37 43 47 47 nf nf
JnehKottifa 0 13 30 37 40 43 43 nf nf
Ψ 136.82*** 74.51*** 43.30*** 28.44*** 17.15*** 492.95*** 497.35***
1)
DMRT
Variety 60.30*** 47.69*** 41.98*** 39.95*** 39.58*** 6.01*** 6.54***
1) Duncan’s multiple range test (DMRT) was conducted for each day among water potentials (Ψ) and varieties
2) nf:
no further germination
3) Level of significance: * p<0.05; *** p<0.001
828 AFEF OTHMANI ET AL.

Table 2. Analysis of variance (F value) of mean day germination (MDG), daily germination speed (DGS) and
germination rate (GR) of 11 durum wheat cultivars for six water potentials.
Variance
Df MDG DGS GR
Sources
Water potential 5 3.06* 1.93 14.86***
Cultivar 10 10.93*** 3.91*** 12.03***
Water potential× cultivar 50 1.13 0.71 0.96
Level of significance: * p<0.05; ***p<0.001

Table 3. Mean day germination (MDG), daily germination speed (DGS) and germination rate (GR) of 11
durum wheat cultivars for six water potentials.
Source of variance
MDG DGS GR
0 0.09 b 1.81 a 2.06 a
-0.47 0.36 a 1.21 ab 1.39 bc
-1.48 0.30 a 1.75 ab 1.30 c
Water potential (bars)
-3.02 0.36 a 1.12 b 1.66 b
-5.11 0.39 a 1.21 ab 2.15 a
-7.73 0.28 a 1.37 ab 1.21 c
Mâali 0.00 d 1.16 c 1.50 bc
Mahmoudi 0.55 a 1.00 c 1.88 ab
Om rabiaa 0.66 a 1.00 c 2.22 a
Karim 0.44 ab 1.00 c 1.88 ab
Nasr 0.61 a 1.00 c 2.16 a
Varieties Salim 0.61 a 1.50 bc 2.00 a
Maghrbi 0.27 bc 1.11 c 1.94 a
Ben bechir 0.11 cd 1.16 c 1.27 cd
Souri 0.00 d 1.72 bc 0.83 e
Agiliglabre 0.00 d 2.27 ab 1.27 cd
Jnehkottifa 0.00 d 2.70 a 0.94 de
Means with similar letter(s) in each trait are not significantly different at p<0.05 (Duncan’s multiple range test)

Table 4. Analysis of variance (F value) of shoot and root-related characters of 11 durum

wheat cultivars for six water potentials.
Variance
df CL (cm) SL (cm) RL (cm) R/S length ratio RN
Sources
*** *** *** ***
Water potential 5 374.74 420.17 165.54 84.27 134.94***
Cultivar 10 49.76*** 18.68*** 9.98*** 5.25*** 3.39***
Water potential× cultivar 50 8.37*** 3.85*** 2.77*** 1.32 4.05***
CL, coleoptile length; SL, shoot length; RL, root length; R/S, root/shoot length ratio; RN, root number

Table 5. The response of five growth parameters of 11 durum wheat cultivars to six water potentials.
Variance sources CL (cm) SL (cm) RL (cm) R/S length ratio RN
Water potential 0 3.38 a 10.44b 11.06 a 1.30 a 5.59 a
(bars) -0.47 3.49 a 11.17 a 10.31 a 0.93b 5.47 ab
-1.48 3.42 a 10.59 ab 10.61 a 1.01b 5.72 a
-3.02 3.31 a 8.66c 6.68c 0.64c 5.41b
-5.11 2.49b 4.78d 7.46b 1.22 a 5.01c
-7.73 0.54c 0.00e 2.45d 0.00d 2.78d
Cultivars Mâali 2.45de 6.86def 7.90c 0.96b 5.21 abc
Mahmoudi 4.20 a 10.45 a 9.93 a 0.77bcd 4.76de
Om rabiaa 2.72c 7.16cde 8.23c 0.93b 5.01bcde
Karim 2.59cd 7.33cde 7.98c 0.89bc 4.94bcde
Nasr 2.10f 6.12f 7.98c 0.93b 5.27 ab
Salim 2.43de 6.47ef 7.59c 0.77bcd 4.62e
Maghrbi 2.28ef 6.92def 9.38 ab 1.19 a 4.85cde
Ben bechir 2.60cd 7.42cd 8.58bc 0.93b 5.08 abcd
Souri 2.95b 9.24b 8.30c 0.70cd 5.43 a
Agiliglabre 2.99b 7.94c 7.56c 0.70cd 4.85cde
Jnehkottifa 3.14b 7.76cd 5.63d 0.61d 4.89bcde
Means with similar letter(s) in each trait are not significantly different at p<0.05 (Duncan’s multiple range test). CL, coleoptile
length; SL, shoot length; RL, root length; R/S, root/shoot length ratio; RN, root number
PEG-INDUCED DOUGHT STRESS TOLERANCE OF DURUM WHEAT 829

Table 6. Interaction effect of osmotic stress and cultivars on Shoot length, root length, root/shoot length ratio and root
seedling traits in 11 durum wheat cultivars. number: The results of ANOVA for SL, RL, R/S length
Water potential
Cultivars CL SL RL RN ratio and RN are described in Table 4. For all these traits
(bars)
0 2.54 9.04 11.41 6.07 there were highly significant differences among water
-0.47 3.11 10.08 8.99 5.60 potentials and among cultivars. Except for R/S length ratio,
-1.48 3.25 8.80 10.67 5.80 there was a significant interaction between osmotic stress
Mâali
-3.02 3.06 8.04 6.54 5.60 levels and cultivars (Table 5). Water deficit significantly
-5.11 2.40 5.22 7.52 5.13 affected root-related traits. Adda et al., (2005) and Faisal et
-7.73 0.39 0.00 2.28 3.07
0 5.21 15.49 13.99 5.27 al., (2017) also showed a significant effect of water stress on
-0.47 5.34 14.66 11.69 5.00 durum and bread wheat root characters.
-1.48 5.51 13.20 14.64 5.87 Early and rapid elongation of roots is an important
Mahmoudi
-3.02 5.65 11.92 8.91 5.27 indicator of drought resistance (El Siddig et al., 2013). RL,
-5.11 2.59 7.48 7.08 5.00 SL and RN decreased as water potential increased (Table 4).
-7.73 0.95 0.00 3.30 2.20
0 3.78 10.63 11.13 5.67
Rana et al., (2017) reported a decrease in RL and SL, which
-0.47 3.61 11.21 11.05 5.47 was an obstacle to cell division in shoot and root elongation,
-1.48 3.57 9.67 10.73 5.47 and to seed reserve utilization. RL increased in Nasr at -0.47
Om rabiaa
-3.02 3.31 8.47 7.89 5.47 bars, in Maâli, Mahmoudi, Souri and Maghrbi at -1.48 bars,
-5.11 1.73 2.99 6.87 5.40 and in Salim under both osmotic stress levels, -0.47 and -
-7.73 0.34 0.00 1.75 2.60
1.48 bars. Baloch et al. (2012) found elongated roots in
0 3.48 10.19 11.20 6.00
-0.47 3.47 11.24 10.17 5.00 wheat cultivars under drought stress. Except for Om rabiaa,
-1.48 3.45 9.73 9.34 5.87 low and moderate osmotic stress (> -7.73 bars) enhanced RN
Karim
-3.02 3.32 9.39 7.98 5.53 in different cultivars. Moreover, most cultivars displayed an
-5.11 1.65 3.45 7.09 5.20 increase in SL when PEG was applied at 50 and 100 g/L,
-7.73 0.18 0.00 2.16 2.07 which might be due to an increase in RL. Compared to
0 2.89 9.55 10.29 6.93
-0.47 2.92 10.38 11.74 6.47
shoots, the physiological activities of roots are less sensitive
-1.48 2.93 9.33 9.35 7.07 to water deficit (Ahmad et al., 2017).
Nasr The varying response of genotypes to PEG treatment,
-3.02 2.41 6.45 7.21 5.80
-5.11 1.23 1.03 7.35 3.80 due to differential genetic sensitivity to water deficit, is very
-7.73 0.22 0.00 1.97 1.60 important for plant breeders as drought-tolerant genotypes
0 3.25 10.97 8.15 5.13 can be screened and tagged at the seedling stage without
-0.47 3.28 10.93 11.56 5.13
-1.48 3.33 9.47 11.02 5.73 extensive and expensive field trials (Meher et al., 2017; Rana
Salim et al., 2017). Durum wheat genotypes forming longer roots
-3.02 2.79 6.36 8.21 5.73
-5.11 1.77 1.10 5.49 4.53 under water limitation exhibit an adaptive response by
-7.73 0.18 0.00 1.14 1.47 increasing water uptake capacity by seeds. There is little
0 2.72 9.27 12.36 5.27 evidence to support that drought-tolerant cultivars uniformly
-0.47 3.06 9.30 11.74 5.60
-1.48 3.06 10.84 12.55 5.40 display advantageous traits such denser shoot and root dry
Maghrbi matter, and longer shoot and root lengths under water stress
-3.02 2.73 7.82 8.93 4.80
-5.11 2.04 4.35 9.17 4.73 (Bin Abdul Hamid, 2012).
-7.73 0.11 0.00 1.54 3.36 Many plants respond to drought by increasing the
0 2.87 9.41 12.35 5.20 proportion of assimilate diverted to growth and thus, increase
-0.47 3.04 9.76 11.50 5.13
-1.48 2.77 10.71 12.25 5.53
the S/R ratio and the volume of soil water available to the
Ben bechir plant (Riaz et al., 2013; Ahmad et al., 2017). The simple
-3.02 2.66 9.33 6.14 5.33
-5.11 3.79 5.37 7.03 5.33 supposition is that deeper and more abundant root systems
-7.73 0.48 0.00 2.24 3.95 can tap extra water from the soil profile, thus moderatingthe
0 3.59 9.95 10.67 6.20 effects of drought (Vadez, 2014). Based on means values of
-0.47 3.64 11.89 8.93 6.20 all cultivars (Table 5), the R/S length ratio was high in the
-1.48 3.69 12.85 10.77 6.23
Souri control (0 bars) with no significant differences in length
-3.02 3.19 8.73 4.65 5.32
-5.11 2.46 6.07 4.34 5.16 under stress (-5.11 bars) and there was no interaction
-7.73 1.18 0.00 3.79 3.53 between water potential and cultivars (Table 6). The higher
0 3.46 10.91 11.07 5.49 R/S length ratio under water stress may be attributed to
-0.47 3.77 12.29 9.80 5.93 longer roots under stress, probably due to the induction of
-1.48 3.08 10.49 8.78 5.36 root-to-shoot hormonal signaling while the root system is
Agiliglabre
-3.02 3.86 9.52 4.31 4.87
-5.11 2.85 4.43 7.13 4.76
subjected to drought stress (Bin Abdul Hamid, 2012). It
-7.73 0.97 0.00 4.32 2.72 could also be associated with higher dry matter and soluble
0 3.41 9.52 9.08 4.28 sugar content in roots, due to an increase in enzyme activity
-0.47 3.16 11.20 6.32 4.67 (Xu et al., 2015).
-1.48 3.01 11.45 6.64 4.60 Mahmoudiformed the longest shoots and roots but had a
Jnehkottifa
-3.02 3.43 9.30 2.79 5.78 low R/S length ratio and RN (Table 5). Root traits associated
-5.11 4.91 5.11 6.41 6.07
-7.73 0.94 0.00 2.57 4.00 with sustaining plant productivity under drought include
CL, coleoptile length; SL, shoot length; RL, root length; RN, small roots with fine diameters, long roots, and dense roots
root number (Comas et al., 2013).
830 AFEF OTHMANI ET AL.

Table 7. Correlation coefficients among germination and seedling characters of 11 wheat cultivars under water stress.
CL (cm) SL (cm) RL (cm) R/S NR MDG DGS GR
LC (cm) 1 0.83** 0.67** 0.44** 0.67** 0.16* -0.12 -0.29**
SL (cm) 1 0.74** 0.46** 0.67** 0.14* -0.07 -0.40**
RL (cm) 1 0.73** 0.64** 0.31** -0.25** -0.25**
R/S 1 0.58** 0.31** -0.23** -0.09
NR 1 0.24** -0.22** -0.27**
MDI 1 -0.77** 0.65**
DGS 1 -0.57**
GR 1
**correlation is significant at p<0.001; * correlation is significant at p<0.05. CL, coleoptile length; SL, shoot length; RL, root length;
R/S, root/shoot length ratio; RN, root number; MDG, mean day germination; DGS, daily germination speed; GR, germination rate

Group I

Group II

Group III

Fig. 1. Biplot of principal component analysis of 11 durum wheat cultivars and studied traits. 1: Maâli, 2: Mahmoudi, 3: Om rabiaa, 4: Karim,
5: Nasr, 6: Salim, 7: Maghrbi, 8: Ben bechir, 9: Souri, 10: Agiliglabre, 11: Jnehkottifa. CL, coleoptile length; SL, shoot length; RL, root
length; R/S, root/shoot length ratio; RN, root number; MDG, mean day germination; DGS, daily germination speed; GR, germination rate.

Relationship between germination and seedling morphological traits such as plant height and 1000-
growth characters kernel weight of 114 durum wheat cultivars derived
from a field-grown trial (Nagel et al., 2014). SL
Correlation analysis: There was a positive and showed a positive and highly significant correlation
significant correlation between MDG and CL, SL, RL with CL (r = 0.83), RL (r = 0.74), R/S length ratio (r =
and R/S length ratio (Table 7). However, a negative and 0.46) and RN (r = 0.67). In addition, RL showed a
highly significant correlation was observed between positive correlation with CL (r = 0.67). Khan et al.
DGS and RL, R/S length ratio and RN. GR showed a (2013) also noted that RL was significantly correlated
negative and highly significant correlation with CL, with CL (r = 0.82) in wheat. The highest correlations
SL, RL, RN and DGS, and a positive correlation with between SL and CL (r = 0.83) and SL and RL (r = 0.74)
MDG. The correlation between GR and R/S length may suggest that selection for these characters can be
ratio was not significant. Rauf et al., (2007) found a useful in breeding programs. Similar results were
significant and positive correlation between GR, CL, obtained by Hellal et al., (2018) in barley cultivars, for
SL and RL but a non-significant and negative three germination periods (3, 5 and 7 days), in which
correlation between GR and R/S length ratio in 16 SL and RL were highly correlated with germination
wheat cultivars. In fact, PEG-induced coleoptile period (r = 0.820, r = 0.829, r = 0.886, and r = 0.871, r
growth under osmotic stress was correlated with agro- = 0.919, r = 0.968).
PEG-INDUCED DOUGHT STRESS TOLERANCE OF DURUM WHEAT 831

Principal component analysis: The relationships Beheshtizadeh, H., A. Rezaie, A. Rezaie and A. Ghandi. 2013.
between different studied traits and cultivars are Principal component analysis and determination of the
graphically presented as a PCA analysis (Fig. 1), as it is selection criteria in bread wheat (Triticum aestivum L.)
the most suitable multivariate method (Beheshtizadeh et genotypes. Int. J. Agric. Crop Sci., 5(18): 2024-2027.
Bin Abdul Hamid, S. 2012. Studies of drought tolerance of hard
al., 2013). The first two components PCA1 (strongly
red winter wheat (Triticum aestivum L.) cultivars in
associated to DGS) and PCA 2 (linked to CL and SL) Nebraska. Thesis, paper 56, Nebraska, Lincoln, USA, 75 pp.
accounted for 77.2% of total variation. Mahmoudi (group Brini, F., M. Hanin, V. Lumbreras, S. Irar, M. Pagès and K.
I) was distinguished from other cultivars, showing Masmoudi. 2007. Functional characterization of DHN-5, a
greatest performance for SL, CL and RL. The latter (RL) dehydrin showing a differential phosphorylation pattern in
was highly correlated to MDS and GR, as assessed by the two Tunisian durum wheat (Triticum durum Desf.) varieties
acute angle of their vectors. Mahmoudi is thus the most with marked differences in salt and drought tolerance.
drought-tolerant cultivar for germination and seedling Plant Sci., 172: 20-28.
traits. Souri, Agiliglabre and Jnehkottifa, which formed Castilhos, G., F. Lazzarotto, L. Spagnolo-Fonini, M.H.
group II, had highest DGS, implying greater germination Bodanese-Zanettini and M. Margis-Pinheiro. 2014.
Possible roles of basic helix-loop-helix transcription factors
ability. Group III includes Maâli, Mahmoudi, Om rabiaa,
in adaptation to drought. Plant Sci., 223: 1-7.
Karim, Nasr, Salim, Maghrbi and Ben bechir, which Chachar, M., N. Chachar, S. Chachar, Q. Chachar, S. Mujtaba
showed a variable response to drought stress. and A. Yousafzai. 2014. In vitro screening technique for
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10(6): 1439-1450.
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most important stages of a plant life cycle and are most Engineering plants to tolerate abiotic stresses. Biocatal.
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(Received for publication 11 July 2018)

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