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because of large volume of soil. In a preliminary study, photometer (Jenway PFP-7). For chloride (Cl-1)
Eucalyptus camaldulensis performed better than many determination, plant samples were extracted with
other eucalyptus species under salt stress in solution HNO3 and chloride was determined from this extract
culture. The present experiment was conducted in a using chloride analyzer (Corning Chloride Analyzer
lysimeter for 22 weeks to study growth performance of 926). The data were analyzed statistically following the
Eucalyptus camaldulensis under salinity and methods of Gomez and Gomez (1984) using MStat-C
waterlogging stresses. (Michigan State University, 1996). The significance of
differences among the means was compared using
Materials and Methods standard error computed as s/√n, where s is the
The experiment was conducted in a rain-protected standard deviation and n shows the number of
wire house at the Institute of Soil & Environmental observations.
Sciences, University of Agriculture, Faisalabad. The
Results and Discussion
average day and night temperatures during the study
were 32 and 20 °C, respectively. Growth parameters
The study was conducted twice in specially Salinity and waterlogging stresses significantly
designed square shaped concrete lysimeters (lxl m2 ) reduced all of the growth parameters significantly
having 30 cm layer of coarse sand at bottom, a thin (Figures 1-4). In most plants growth gradually reduces
layer of glass wool in between sand and a soil column as salinity increases above a threshold value which
of 133 cm. The total profile (sand + soil) was of 165 varies in different species (Saqib et al., 2004; 2005;
cm. Three salinity levels used in the study were: S1 Tahir et al., 2006). Mean tree height of Eucalyptus
(control: 3.5 dS m-1), S2 (15 dS m-1) and S3 (30 dS m-1). camaldulensis decreased significantly as salinity
Salinity was developed by saturating the soil with increased in the root medium at waterlogging as well as
salinized water (NaCl was dissolved in calculated non-waterlogging treatment (Figure 1). Similar
amount of canal water). Half amount of water was reduction in tree height was observed in Eucalyptus in a
applied from surface and the other half from bottom of solution culture experiment (Nasim et al., 2008). Effect
the lysimeter through provisions of inlet and outlet of waterlogging on tree height was non-significant at
holes. control (3.5 d Sm-1) and S2 treatment (15 d Sm-1)
After attaining desired salinity levels, six uniform however, waterlogging significantly depressed tree
6-month-old saplings of Eucalyptus camaldulensis height at S3 (30 d Sm-1). The reduction in mean tree
(Local) were transplanted in each Lysimeter. Six weeks height was 1.5% due to waterlogging, 7.6% due to S2,
after transplanting, plants were thinned to 4. At this 10% due to combined effect of S2 and waterlogging,
stage waterlogging/anaerobic stress (in half of the 45.5% due to S3 and 59.2% due to combined effect of
lysimeters) was imposed and maintained continuously S3 and waterlogging.
for 12 weeks. However, soil remained in saturated 250
150
was measured and numbers of branches per plant were
counted. Leaves from old (the lower most two
branches from base of the tree), middle (branches 100
S1, while it was lowest at S3 (EC 30 d Sm-1) both at (waterlogging alone) to 92.2% (waterlogging with 30 d
waterlogging and non-waterlogging conditions. Sm-1salinity). Many of earlier scientists also reported
Waterlogging alone did not affect NB at control significant reduction in biomass under salt stress in
treatment, however, it reduced NB when combined with eucalyptus (Qureshi et al., 1993; Marcar et al., 1995).
S2 (15 d Sm-1) and S3 (30 d Sm-1) treatment (Figure 2). 400
Number of branches was about 47% lower at S3 and
waterlogging treatment compared to only S3. 350 Control
Water Logging
F re s h w e ig h t o f tw ig s (g /p la n t)
50 300
45 Control
250
40 Water Logging
200
35
N u m b e r o f b ra n c h e s
30 150
25
100
20
50
15
10 0
Control 15 d Sm-1 30 d Sm-1
5
Salinity levels
0
Control 15 d Sm-1 30 d Sm-1 Fig. 3. Effect of salinity and waterlogging on fresh weight of twigs of Eucalyptus
camaldulensis
Salinity Levels
400
Fig. 2. Effect of salinity and water logging on number of branches of Eucalyptus
camaldulensis Control
350
Water Logging
There was significant main and interactive effect of
F re s h w e ig h t o f le a v e s (g /p la n t)
300
aggravated significantly as plants were affected by Fig. 4. Effect of salinty and waterlogging on fresh weight of leaves of
Eucalyptus camaldulensis
waterlogging. The relative reduction in fresh weight of
twigs was 33.9%, 56.8%, 79.2% and 89.5% in the case
of S2 (15 d Sm-1) alone, S2 (15 d Sm-1) and Physiological parameters
waterlogging, S3 (30 d Sm-1) alone and S3 and
waterlogging, respectively. Tissue ion concentrations are the result of ion
transport and growth of shoot. The balance between
There was significant main and interactive effect water and salt uptake could be maintained by reducing
on fresh weight of leaves (FWL) of Eucalyptus (Figure transpiration but at the expense of reduced carbon
4). Fresh weight of leaves of Eucalyptus camaldulensis fixation and reduced growth rate (Marschner, 1995).
was significantly reduced as salinity was increased in The rate of ion transport depends not only on the rate of
the root medium. Fresh weight of leaves was about 75% shoot growth but also on the external concentration of
and 30% at S2 (15 d Sm-1) and S3 (30 d Sm-1) levels, salts. Salt stress affects uptake, transport and utilization
respectively as compared to non-waterlogging control. of different nutrients (Marschner, 1995; Grattan and
Waterlogging alone did not affect FWL of eucalyptus; Grieve, 1999; Zhu, 2003). The imbalance in nutrient
however, it aggravated the reduction in FWL due to uptake under salinity stress may result in excessive
salinity at 15 d Sm-1 and 30 d Sm-1. The minimum fresh accumulation of Na+ and Cl- in tissue (Saqib et al.,
weight of leaves was recorded under the combined 2004; Tahir et al., 2006) and ultimately reduction in
stress of high salinity + waterlogging. The relative crop yield. Hence concentration of Na+ in plants is a
reduction in fresh weight of leaves varied from 2.2% good indicator of salinity tolerance.
A lysimeter study of Eucalyptus camaldulensis under salinity and waterlogging stress 95
There was significant main and interactive affect of significantly (p<0.05) in leaves of Eucalyptus
salinity and waterlogging on Na+ concentration in camaldulensis compared with control and waterlogging
lower, middle and upper leaves of Eucalyptus treatment. Waterlogging alone did not have an effect on
camaldulensis (Table 1). Increased salinity in the root Cl- concentration, but combined stress of salinity
medium, significantly increased Na+ concentration (>2 andwaterlogging resulted in marked increase in Cl-
folds) in leaves of Eucalyptus compared to control concentration (> 2 folds). The highest Cl- concentration
(Table 1). Na+ was significantly lower in young leaves was observed under S3 alone and S3 and waterlogging.
at all salinity treatments at non-waterlogging Lowest leaf Cl- concentration was found in control and
conditions. Waterlogging alone did not increase mean waterlogging alone Saqib et al. (2004) also reported an
Na+ concentration in leaves as compared to control, but increase in Cl- concentration under salinity stress.
waterlogging + salinity resulted in a significant increase Chloride concentration was significantly lower in upper
(2.5-4.5 folds) in the Na+ concentration of leaves from leaves at all treatments compared to middle and lower
different branches. The accumulation of Na+ was the leaves.
highest (632 mmole Na kg-1) at S3 and waterlogging
Adequate K+ concentration in plant tissues is
followed by S2 and waterlogging (321 mmole Na kg-1)
essential for survival particularly in saline soils
and S3 (295 mmol kg-1). Interactive effect of salinity X
(Marschner, 1995). High concentration of Na+ in root
waterlogging clearly showed that accumulation of Na+
environment can restrict K+ acquisition (Subbarao et
in leaves increased when the plants were exposed to
al., 1990; Liu et al., 2001; Saqib et al., 2004) leading to
dual stress of waterlogging and salinity (30 d Sm-1).
inhibition of K requiring processes in the cytoplasm.
Na+ in lower leaves was maximum at all salinity
Nutritional imbalances as a growth limiting factor
treatment compared to other leaves. Saqib et al. (2004)
under saline and /or waterlogging conditions has also
and Tahir et al. (2006) also reported significant increase
been stressed by Grattan and Grieve (1999) and Tahir et
in Na+ concentration in leaves of different wheat
al. (2006). Reduced K concentration in leaves under
cultivars under salinity stress and reported a significant
salinity stress in present study also revealed that poor
reduction in growth of plants with an increase in Na+
growth of plants in saline and waterlogging condition
concentration in leaves. In our preliminary solution
may not only due to decreased water potential (osmotic
culture experiments, different eucalyptus species
effect) but also due to ionic imbalances particularly of
accumulated higher amounts of Na in their leaves under
Na+,Cl- and K+. There was significant main and
salinity treatment (Nasim et al., 2008).
interactive effect of salinity and waterlogging on K+
There were significant main and interactive effect concentration in leaves of eucalyptus (Table 2).
of salinity and waterlogging on C1- concentration in
leaves (Table 1). Salinity enhanced C1- accumulation
Table 1. Leaf sodium (Na+) and chloride (Cl-) concentration in Eucalyptus camaldulensis grown with salinity
and waterlogging
Parameters Branches Non-waterlogging Waterlogging
Table 2. Leaf potassium (K+) and K+:Na+ ratio in Eucalyptus camaldulensis grown with salinity and
waterlogging
Parameters Branches Non-waterlogging Waterlogging
Overall treatment comparison revealed that Seminar on Prospects of Saline Agriculture in the
waterlogging alone and as well as along with high GCC Countries, March, 18-20, 2001. Dubai, UAE.
salinity significantly (p<0.01) depressed K+ Ghafoor, A., M. Qadir and G. Murtaza. 2004. Salt
accumulation in leaves compared with control. Saqib et Affected Soils: Principles of Management. Allied
al. (2004) and Tahir et al. (2004) also reported Book Centre, Lahore.
significant reduction in K concentration in leaves of Grattan, S.R. and C.M. Grieve. 1999. Salinity-mineral
plants grown with salinity in hydroponics experiments. nutrient relations in horticultural crops. Scientia
Significant differences were found in the concentration Horticulturae 78: 127-157.
of K+ in leaves belonging to different aged branches. Jackson, M.B. 1979. Rapid injury to peas by soil water
The leaves of young branches maintained high logging. Journal of the Science of Food and
concentration of K+ whereas it was the poorest in the Agriculture 30: 143-152.
case of leaves on old branches. Eucalyptus Liu, W., D.J. Fairbairn, R.J. Reid and D.P. Schachtman.
camaldulensis (Local) accumulated less Na+ and Cl- 2001. Characterization of two HKT1 homologues
and more K+ in young leaves than older ones. One from Eucalyptus camaldulensis that display
adaptation to salinity and other stresses including intrinsic osmosensing capability. Plant Physiology
waterlogging in plants, is to rely on the youngest leaves 127: 283-294.
for photosynthetic CO2 assimilation. Marcar, N.E., D.F. Crawford, P.M. Leppert, T.
Jovanovic, R. Floyd and R. Rarrow. 1995. Trees
Conclusion for salt land: A guide for selecting native species
Salinity significantly depressed all of the growth for Australia. CSIRO, Australia.
parameters in E. camaldulensis. Waterlogging alone did Marschner, H. 1995. Mineral Nutrition of Higher
not affected growth parameters; however in Plants. Academic Press, London.
combination with salinity it significantly affected Morard, P. and J. Silvestre. 1996. Plant injury due to
growth parameters and ionic composition of leaves of oxygen deficiency in the root environment of soil
E. camaldulensis (local). Na+ and Cl- concentration was less culture: a review. Plant and Soil 184: 243-254.
significantly more in old leaves indicating an adoption Munns, R. 2005. Genes and salt tolerance: bringing
of E. camaldulensis against salinity stress. Na+ and Cl- them together. New Phytologist 167: 645-663.
concentration in leaves were significantly increased Nasim, M., R.H. Qureshi, T. Aziz, M. Saqib, S. Nawaz,
when plants were subjected to dual stress of salinity and S.T. Sahi and S. Pervaiz. 2008. Growth and ionic
waterlogging compared to salinity and waterlogging composition of salt-stressed Eucalyptus
alone. camaldulensis and Eucalyptus tereticornis.
Pakistan Journal of Botany 40(2): 799-805.
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