Documente Academic
Documente Profesional
Documente Cultură
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
To link to this Article: DOI: 10.1080/01904160903092648
URL: http://dx.doi.org/10.1080/01904160903092648
This article may be used for research, teaching and private study purposes. Any substantial or
systematic reproduction, re-distribution, re-selling, loan or sub-licensing, systematic supply or
distribution in any form to anyone is expressly forbidden.
The publisher does not give any warranty express or implied or make any representation that the contents
will be complete or accurate or up to date. The accuracy of any instructions, formulae and drug doses
should be independently verified with primary sources. The publisher shall not be liable for any loss,
actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directly
or indirectly in connection with or arising out of the use of this material.
Journal of Plant Nutrition, 32: 1427–1439, 2009
Copyright © Taylor & Francis Group, LLC
ISSN: 0190-4167 print / 1532-4087 online
DOI: 10.1080/01904160903092648
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.
INTRODUCTION
1427
1428 M. Nasim et al.
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.
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.
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.
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
1430 M. Nasim et al.
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).
Downloaded By: [Nasim, Muhammad] At: 06:19 6 August 2009
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
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).
Salinity × Hypoxia Interaction in Eucalyptus 1431
Downloaded By: [Nasim, Muhammad] At: 06:19 6 August 2009
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.
1432 M. Nasim et al.
Downloaded By: [Nasim, Muhammad] At: 06:19 6 August 2009
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.
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.
DISCUSSION
Table 1
Relationship between different growth parameters and ionic composition of young and
mature leaves of Eucalyptus species
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
Sodium uptake
E. tereticornis 3.8 3.2
E. camaldulensis (Silverton) 2.9 2.8
E. camaldulensis (Local) 1.4 2.3
Chloride uptake
E. tereticornis 1.6 2.0
E. camaldulensis (Silverton) 1.3 1.4
E. camaldulensis (Local) 1.1 1.7
Potassium uptake
E. tereticornis 0.95 0.76
E. camaldulensis (Silverton) 1.0 0.69
E. camaldulensis (Local) 1.0 1.4
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.
1436 M. Nasim et al.
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-
Downloaded By: [Nasim, Muhammad] At: 06:19 6 August 2009
REFERENCES