Salinty N Hypoxia Interaction in Eucalptus

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Different Eucalyptus Species Show Different Mechanisms of Tolerance to

Salinity and Salinity × Hypoxia
Muhammad Nasim a; Riaz H. Qureshi b; Tariq Aziz a; M. Saqib b; Shafqat Nawaz a; J. Akhtar b; M. A. Haq b;
Shahbaz Talib Sahi a
a
Sub-Campus Depalpur, University of Agriculture, Faisalabad, Depalpur, Pakistan b Saline Agriculture
Research Centre, University of Agriculture, Faisalabad, Depalpur, Pakistan
Online Publication Date: 01 September 2009
To cite this Article Nasim, Muhammad, Qureshi, Riaz H., Aziz, Tariq, Saqib, M., Nawaz, Shafqat, Akhtar, J., Haq, M. A. and Sahi,
Shahbaz Talib(2009)'Different Eucalyptus Species Show Different Mechanisms of Tolerance to Salinity and Salinity ×
Hypoxia',Journal of Plant Nutrition,32:9,1427 — 1439
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Journal of Plant Nutrition, 32: 1427–1439, 2009
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ISSN: 0190-4167 print / 1532-4087 online
DOI: 10.1080/01904160903092648
Different Eucalyptus Species Show Different

Mechanisms of Tolerance to Salinity
and Salinity × Hypoxia
Muhammad Nasim,1 Riaz H. Qureshi,2 Tariq Aziz,1 M. Saqib,2

Shafqat Nawaz,1 J. Akhtar,2 M. A. Haq,2 and Shahbaz Talib Sahi1
1
Sub-Campus Depalpur, University of Agriculture, Faisalabad,
Downloaded By: [Nasim, Muhammad] At: 06:19 6 August 2009
Depalpur, Pakistan
2
Saline Agriculture Research Centre, University of Agriculture, Faisalabad,
Depalpur, Pakistan
ABSTRACT
We studied the effect of sodium chloride (NaCl) salinity and oxygen deficiency stress
on growth and leaf ionic composition of three Eucalyptus species [E. tereticornis, E.
camaldulensis (Silverton), and E. camaldulensis (Local)]. Species were grown with
control (no NaCl) and salinity (150 mol m−3 NaCl) under hypoxic and non-hypoxic
conditions in nutrient solution with five replications following CRD. Species differed
significantly in their response to salinity and hypoxia. Absolute shoot dry matter was
significantly better in E. camaldulensis (Silverton) in salinity and in E. camaldulensis
(Local) in saline hypoxic treatment. E. tereticornis was the most sensitive species to
salinity and salinity × hypoxia in the root environment. Sodium (Na+ ) and chloride (Cl− )
concentrations were significantly lower in E. camaldulensis (Local) in non-hypoxic
saline treatment compared to the other two species. E. camaldulensis (Silverton) seems
to have better tissue compartmentalization, whereas E. camaldulensis (local) seems to
have better exclusion of Na+ at the root level.
Keywords: salinity, hypoxia, Eucalyptus, leaf expansion, growth, ionic composition
INTRODUCTION
Desertification of arable lands is a serious threat to agriculture around the globe.

Salinization is an important factor contributing to the degradation of the arable
Received 29 October 2007; accepted 17 July 2008.

Address correspondence to Dr. Muhammad Nasim, University of Agriculture,
Faisalabad, Sub-Campus Depalpur, Depalpur, Okara, Pakistan. E-mail: mnasimshahid@
yahoo.com
1427
1428 M. Nasim et al.
lands particularly in arid and semi-arid regions (Ashraf, 1994). Approximately

6.3 million hectares of agricultural land in Pakistan is affected by varying
degrees of salinity/sodicity (Ghafoor et al., 2004) that adversely affects plant
growth and yield in a number of species (Eom et al., 2007). Soil salinity affects
plant growth mainly through osmotic effects and specific ion toxicities (Grattan
and Grieve, 1999; Munns, 2002; 2005) resulting from accumulation of sodium
and chloride in the plant tissues (Saqib et al., 2004a; 2005a; Rezaei et al.,
2006). Sodium (Na) concentration in tissues and potassium (K+ ): Na+ ratios
are extensively used in screening mass germplasms as selection parameters
(Saqib et al., 2004a; Tahir et al., 2006).
Salt affected soils (mainly those with high sodium) are generally dispersed
soils having low soil permeability, porosity, and hydraulic conductivity (Qadir
et al., 2005). Low water infiltration and soil dispersion poses a serious drainage
problem and may lead to waterlogging. Plant roots affected by waterlogging are
deprived of sufficient oxygen resulting in a change in the mode of respiration
from aerobic to an-aerobic and low energy production (Marschner, 1995). Low
energy production in roots disturbs the nutrient and water uptake (Jackson,
1979; Morard and Silvestre, 1996). Waterlogging also reduces nutrient uptake
that results in nutrient deficiency in shoot leading to reduced shoot growth
(Trought and Drew, 1980). Higher abscisic acid production and movement
to younger leaves under oxygen (O2 ) deficiency causes stomatal closure in
plants (Zhang and Zhang, 1994) affecting photosynthesis and ultimately yield.
Waterlogging also affects sodium exclusion from plant roots, which is a major
salinity tolerance mechanism in various glycophytes (Barrett-Lennard, 1986;
Saqib et al. 2005a). A combined effect of salinity and waterlogging on plant
growth is more damaging than caused by salinity and waterlogging alone
(Qureshi and Barrett-Lennard, 1998; Saqib et al., 2004a).
Although several relatively salt tolerant cultivars of different agronomic
crops are available to grow on moderately salt affected soils yet these culti-
vars fail to produce economical yields on highly degraded salt affected soils.
Reclamation of such soils through physical and chemical approaches is also
not feasible. Revegetation of these lands with salt tolerant tree species is a vi-
able approach mainly because of low input and increased demand of wood for
fuel and furniture. Selection of suitable tree species is a pre-requisite for such
biological remediation approach. However, very little work has been reported
on this aspect particularly on tree species. The present paper reports the effect
of salinity and hypoxia interaction on growth and leaf ionic composition of
three Eucalyptus species.
MATERIALS AND METHODS
Site and Environmental Conditions
The experiment was conducted in a rain-protected wire house at the Insti-

tute of Soil & Environmental Sciences, University of Agriculture, Faisalabad,
Salinity × Hypoxia Interaction in Eucalyptus 1429
Pakistan. Faisalabad is situated at 73.4◦ longitude and 31.5◦ latitude. The aver-
age day and night temperatures during the study were 32 and 20◦ C, respectively,
while the average maximum relative humidity was 70%, and there was 7 h av-
erage daily sunshine during the growth period.
Plant Material and Experimental Details
Seeds of three Eucalyptus species [E. tereticornis, E. camaldulensis (Silver-

ton), and E. camaldulensis (Local)] were collected from Australian Revegeta-
tion Corporation Limited, Western Australia and Saline Agriculture Research
Centre, University of Agriculture, Faisalabad, Pakistan. Fifty seeds of each
species were sown in polyethylene lined iron trays containing silica gravel and
synthetic vermiculite (mixed in 1:1 ratio). The canal water was used to moisten
the seeds for proper germination. Electrical conductivity of canal water was
0.29 d Sm−1 . One week after germination, half strength Hoagland’s nutrient
solution (Hoagland and Arnon, 1950) was used for the seedling establishment.
Seedlings were grown for three months in these trays. Uniform sized seedlings
were then transplanted in foam plugged holes in polystyrene sheets floating
on continuously aerated half strength Hoagland’s nutrient solution contained
in polyethylene lined iron tubs of 200 L capacity (100 × 100 × 30 cm3 ).
The experiment was replicated five times following completely randomized
design (CRD). Solution pH was monitored and maintained daily at 6 ± 0.5.
There were four treatments viz i) non-saline non-hypoxic (control), ii) saline
non-hypoxic, iii) non-saline hypoxic, and iv) saline-hypoxic. Salinity level for
saline treatment was 150 mol m−3 sodium chloride (NaCl). The salinity was
developed in three equal splits in a week (each of 50 mol m−3 NaCl after 2
d). The hypoxia was imposed by surface sealing of the nutrient solution. Three
days after surface sealing, substrate oxygen concentration (measured with O2
electrode) was 3 mg dm−3 . The treatment solutions were replaced thereafter
weekly till harvesting and in hypoxic treatments the new solution was flushed
with N2 for 15 minutes to remove oxygen.
Growth Measurement
To study leaf expansion (increase in leaf length) 2-day old leaf of each plant
from shoot apex was marked and leaf length was measured at 2 day (d) interval
up to 11th day. After 8 weeks of salinity and hypoxia stress plant height was
recorded. At this time plants were harvested and their dry weights of shoots
and roots were recorded after oven drying the samples at 70◦ C in a vacuum
oven for 48 hrs.
Leaf Ionic Analysis
Plants were harvested after 8 weeks and leaves of plants were divided into top
leaves (young leaves) and lower fully expanded leaves (mature leaves). The
separated leaves were immediately washed with distilled water and blotted dry
with tissue paper. The samples were then dried at 70◦ C in a forced air driven
oven for 48 h. The oven dried plant samples were fine ground in a wily mill to
pass through 1 mm sieve. The fine ground plant samples (1 g) were digested in
tri-acid mixture (sulfuric acid, nitric acid, and perchloric acid) (Miller, 1998).
Potassium and Na+ were determined on a flame photometer (Jenway PFP-
7, Bibby Scientific LTD., Essex England). For chloride determination, plant
samples were extracted with HNO3 and chloride was determined from this
extract using chloride analyzer (Corning Chloride Analyzer 926, Corning Inc.,
Corning, NY, USA).
Statistical Analysis
The data were analyzed statistically following the methods of Gomez and
Gomez (1984) using MSTAT-C (Russell and Eisensmith, 1983). The signif-
icance of differences
√ among the means was compared using standard error
computed as s/ n, where s is the standard deviation and n shows the number
of observations.
RESULTS
Shoot and Root Growth
Salinity significantly reduced shoot dry matter of all the species at both non-
hypoxic and hypoxic conditions (Figure 1). E. camaldulensis (Silverton) pro-
duced the maximum absolute and relative shoot dry matter (36%) under non-
hypoxic salinity followed by E. camaldulensis (Local) and E. tereticornis.
However, under hypoxic salinity treatment E. camaldulensis (Local) performed
best followed by E. camaldulensis (Silverton) and E. tereticornis, in absolute
as well as relative terms. Hypoxia alone reduced the shoot and root dry matter
of E. tereticornis only. Root dry matter (RDM) of Eucalyptus species was also
significantly reduced by salinity in hypoxic conditions (Figure 2). E. camal-
dulensis (Local) and E. tereticornis produced respectively, the maximum and
minimum root dry matter in this treatment. However, in non-hypoxic salinity
treatment E. camaldulensis (Silverten) and E. camaldulensis (Local) differed
non-significantly. Root: shoot ratio (RSR) of Eucalyptus species was increased
significantly under saline conditions both in hypoxic and non-hypoxic treat-
ments, whereas hypoxia alone did not affect it significantly (data not shown).
Salinity also significantly reduced leaf expansion of E. tereticornis and E.
camaldulensis (Local) under non-hypoxic as well as hypoxic conditions (Fig-
ure 3). Plant height of all the species was also reduced significantly by salinity
and salinity × hypoxia (data not shown).
Figure 1. Shoot dry matter (g/plant) of Eucalyptus species grown with salinity and
hypoxia. The columns show mean of 5 replications and bars show standard error.
Salinity level for saline treatment was 150 mol m−3 NaCl. Plants were grown for 8
weeks in the treatment solutions.
Figure 2. Root dry matter (g/plant) of Eucalyptus species grown with salinity and
hypoxia. The columns show mean of 5 replications and bars show standard error.
Salinity level for saline treatment was 150 mol m−3 NaCl. Plants were grown for 8
weeks in the treatment solutions.
Figure 3. Leaf expansion (cm/d) of Eucalyptus species grown with salinity and hy-
poxia. The columns show mean of 5 replications and bars show standard error. Salinity
level for saline treatment was 150 mol m−3 NaCl. Plants were grown for 8 weeks in the
treatment solutions.
Leaf Ionic Composition
Leaf ionic composition of young as well as mature leaves was significantly

affected by salinity under non-hypoxic as well as hypoxic conditions with a
similar trend in both types of the leaves (Figures 4–6; data shown only for
the young leaves). Salinity caused a significant increase in Na+ and Cl− con-
centrations in both type of the leaves irrespective of the hypoxic treatment
(Figures 4 and 5). Sodium concentration in young leaves of plants grown with
salinity was 3.5 folds and 4.5 folds higher than those grown without salinity
under non-hypoxic and hypoxic conditions, respectively. E. tereticornis and E.
camaldulensis (Silverton) accumulated the maximum Na+ under non-hypoxic
saline treatment and differed non-significantly. E. camaldulensis (Local) accu-
mulated the minimum leaf Na+ in both the saline treatments. Leaf Na+ was
also significantly higher in the leaves of all the species under hypoxic salinity
treatment than under non-hypoxic salinity treatment. The maximum chloride
(Cl− ) concentration was observed in the leaves of E. camaldulensis (Silverton)
under non-hypoxic salinity and in the leaves of E. tereticornis under hypoxic
salinity treatment. E. camaldulensis (Local) differed non-significantly with E.
camaldulensis (Silverton) under hypoxic saline conditions and with E. tereti-
cornis under non-hypoxic saline conditions. Potassium sodium ratio (K+ : Na+
ratio) was significantly decreased due to salinity stress in the root environment
Figure 4. Sodium concentration (mmoles kg−1 ) in young leaves of Eucalyptus species

grown with salinity and hypoxia. The columns show means of 5 replications and bars
show standard error. Salinity level for saline treatment was 150 mol m−3 NaCl. Plants
were grown for 8 weeks in the treatment solutions.
Figure 5. Chloride concentration (mmoles kg−1 ) in young leaves of Eucalyptus species

grown with salinity and hypoxia. The columns show means of 5 replications and bars
show standard error. Salinity level for saline treatment was 150 mol m−3 NaCl. Plants
were grown for 8 weeks in the treatment solutions.
Figure 6. K:Na in young Leaves of Eucalyptus species grown with salinity and hy-
poxia. The columns show means of 5 replications and bars show standard error. Salinity
level for saline treatment was 150 mol m−3 NaCl. Plants were grown for 8 weeks in the
treatment solutions.
under hypoxic as well as non-hypoxic conditions (Figure 6). The maximum

K+ : Na+ ratio was observed in E. camaldulensis (Local) followed by E. camal-
dulensis (Silverton) under both non-hypoxic and hypoxic salinity treatments.
Salinity under hypoxic conditions also have a significantly higher effect than
under non-hypoxic conditions whereas hypoxia alone significantly decreased
the leaf K+ : Na+ of E. tereticornis only.
DISCUSSION
Sodium chloride salinity in the nutrient solution significantly reduced tree

growth in terms of plant height, leaf expansion, shoot dry matter, and root dry
matter in all the species (Figures 1–3) and there was a negative significant
correlation between growth performance and leaf Na+ and Cl− concentrations
(Table 1). Water stress, ion imbalance and ion toxicity are considered the
common causes of growth reduction due to salinity (Munns, 2002; 2005; Zhu,
2003). Genotypic differences for different growth parameters were significant
among the three species in non-hypoxic saline treatment. In this treatment, E.
camaldulensis (Silverton) and E. camaldulensis (Local) produced higher shoot
and root dry matter than E. tereticornis but differed non-significantly with one
Table 1
Relationship between different growth parameters and ionic composition of young and
mature leaves of Eucalyptus species
Shoot dry matter Root dry matter Leaf expansion
Na+ in young leaves −0.71∗∗ −0.60∗ −0.39NS

K+ in young leaves 0.51∗ 0.72∗ 0.47∗
K+ :Na+ in young leaves 0.82∗∗ 0.79∗∗ 0.61∗
Cl− in young leaves −0.69∗ −0.63∗ −0.47∗
another. The poor growth performance of E. tereticornis may be due to higher

Na+ and Cl− uptake, lower K+ uptake (Table 2) and resultantly low K+ : Na+
ratio in its leaves. The leaf Na+ concentration of E. camaldulensis (Silverton)
was statistically similar to E. tereticornis but its better growth shows better
compartmentalization of Na+ and Cl− into the vacuoles by this species. The
higher K+ : Na+ ratio also supports its higher growth performance (Rezaei et
al., 2006). Saqib et al. (2005b) reported that better compartmentalization of
Na+ into the vacuoles is an important determinant for salt tolerance in wheat.
Lower Na+ and Cl− accumulation by E. Camaldulensis (Local) shows better
exclusion in this species at the root level (Table 2; Na+ and Cl− uptake per g
root dry matter) that enabled its better growth (Saqib et al., 2004a). A number of
Table 2
Sodium and chloride uptake (mg per g root dry matter) by different Eucalyptus
species under saline and saline hypoxic conditions
Non-hypoxic saline Hypoxic saline
Sodium uptake
E. tereticornis 3.8 3.2
E. camaldulensis (Silverton) 2.9 2.8
E. camaldulensis (Local) 1.4 2.3
Chloride uptake
Potassium uptake
The columns show mean of 5 replications and bars show standard error.
Salinity level for saline treatment was 150 mol m−3 NaCl. Plants were grown
for 8 weeks in the treatment solutions.
researchers have reported significant variations in the growth and salt tolerance
among different species of the woody plants (Bell et al., 1994; Marcar et al.,
1995; Rawat and Banerjee, 1998). Many earlier researchers have used Na+ :
K+ ratio as an indicator of salinity tolerance (Saqib et al., 2004b; Munns, 2005;
Rezaei et al., 2006) as adequate Na+ : K+ ratio in the cytosol is essential for
normal cellular functions of plants (Marschner, 1995; Chinnusamy et al., 2005).
Higher levels of external Na+ in saline soils interfere with K+ acquisition by
plants (Subbarao et al., 1990). The species with ability to maintain K+ uptake
or to exclude Na+ can tolerate salinity stress such as E. camaldulensis (local).
The lower Na+ uptake and higher K+ uptake at root level, and higher K+ :
Na+ ratio in this species is in accordance with its better performance at saline
environment (Table 2; Figure 6).
Low oxygen supply in hypoxia affects root growth because energy pro-
duction is decreased (Barret-Lennard, 1986). Under hypoxia, shoot and root

growth retardation has also been reported by several researchers (Jackson,
1979; Lizaso et al., 2001). In the present study, hypoxia significantly reduced
the shoot and root growth of the salt sensitive species E. tereticornis only (Fig-
ures 1-2). Aerenchyma development in the roots is an important mechanism of
hypoxia tolerant (Saqib et al., 2005a) that may not have developed considerably
in this species. Marcar et al. (1995) also observed large variations among the
eucalyptus species for their tolerance to oxygen deficiency (waterlogging). In
saline-hypoxic conditions roots are not able to exclude Na+ and Cl− mainly
because of low energy production (Qureshi and Barret-Lennard, 1998). This
lower ability to exclude toxic elements can further inhibit K+ uptake by plant
roots. Saqib et al. (2004a) reported a higher accumulation of Na+ and Cl− and
lower K+ in leaf sap under saline waterlogged conditions than under saline
conditions. Marcar et al. (1993) and Galloway and Davidson (1993) observed
similar depressive effects of salinity × hypoxia interaction on growth of Euca-
lyptus and Atriplexes.
However, in the present study higher reduction in shoot and root growth
was observed only in E. camaldulensis (Silverton) in saline hypoxic treatment
than in the non-hypoxic saline treatment (Figures 1–2). It may be due to its
decreased root ability to develop aerenchyma that resulted in low Na+ exclusion
and K+ uptake at the root level (Figure 4; Table 2) as aerenchyma helps in root
Na+ exclusion (Saqib et al., 2005b). In this treatment E. camaldulensis (local)
performed significantly better than the other species which may be due to its
better aerenchyma and nodal root development and hence better salt exclusion
at the root level. In conclusion, E. camaldulensis (Silverton) is better tolerant
to salinity alone and E. camaldulensis (local) is better tolerant to saline and
hypoxic conditions. E. camaldulensis (Silverton) seems to have better tissue
compartmentalization whereas E. camaldulensis (local) seems to have better
exclusion of Na+ at the root level.
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Different Eucalyptus Species Show Different Mechanisms of Tolerance to

Online Publication Date: 01 September 2009

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Different Eucalyptus Species Show Different

Muhammad Nasim,1 Riaz H. Qureshi,2 Tariq Aziz,1 M. Saqib,2

Keywords: salinity, hypoxia, Eucalyptus, leaf expansion, growth, ionic composition

Desertification of arable lands is a serious threat to agriculture around the globe.

Received 29 October 2007; accepted 17 July 2008.

lands particularly in arid and semi-arid regions (Ashraf, 1994). Approximately

MATERIALS AND METHODS

Site and Environmental Conditions

The experiment was conducted in a rain-protected wire house at the Insti-

Plant Material and Experimental Details

Seeds of three Eucalyptus species [E. tereticornis, E. camaldulensis (Silver-

Leaf Ionic Analysis

Shoot and Root Growth

Leaf Ionic Composition

Leaf ionic composition of young as well as mature leaves was significantly

Figure 4. Sodium concentration (mmoles kg−1 ) in young leaves of Eucalyptus species

Figure 5. Chloride concentration (mmoles kg−1 ) in young leaves of Eucalyptus species

under hypoxic as well as non-hypoxic conditions (Figure 6). The maximum

Sodium chloride salinity in the nutrient solution significantly reduced tree

Shoot dry matter Root dry matter Leaf expansion

Na+ in young leaves −0.71∗∗ −0.60∗ −0.39NS

another. The poor growth performance of E. tereticornis may be due to higher

Non-hypoxic saline Hypoxic saline

duction is decreased (Barret-Lennard, 1986). Under hypoxia, shoot and root

Ashraf, M. 1994. Breeding for salinity tolerance in plants. Critical Reviews in

proving salt tolerance in plants. Crop Science 45: 437–448.

Miller, R. O. 1998. Nitric-percloric wet digestion in an open vessel. In: Hand-

rigated Land in Pakistan: A Handbook. Canberra, Australia: Australian

Trought, M. C. T., and M. C. Drew. 1980. The development of waterlogging

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