Plant Soil (2017) 418:477–491
DOI 10.1007/s11104-017-3309-7
REGULAR ARTICLE
The effect of salinity on plant-available water
Anna R. Sheldon & Ram C. Dalal & Gunnar Kirchhof &
Peter M. Kopittke & Neal W. Menzies
Received: 10 December 2016 / Accepted: 2 June 2017 / Published online: 17 June 2017
# Springer International Publishing AG 2017
Abstract
Aims Plant-available water is determined by soil matric
and osmotic potential. The effect of salinity is a combi-
nation of the osmotic potential, the plant’s capacity to
osmotically adjust, and the specific toxicity of the salt.
Our aim was to better understand the relative importance
of these components in a soil where the relationship
between soil solution composition and soil water con-
tent had been characterized.
Method The capacity of wheat (Triticum aestivum L.)
and chickpea (Cicer arietinum L.) to extract water from
a saline soil was examined by imposing water stress on
established plants, which were then grown until perma-
nent wilting point (PWP) was reached.
Results Wheat extracted soil moisture to lower poten-
tials (−1.2 MPa) than chickpea (−0.80 MPa) in 0 NaCl
treatments. Where salinity was low to moderate, plants
extracted water to a PWP determined by the combined
total of matric and osmotic potentials. Wheat extracted
water to PWP in salinity treatments producing saturated-
paste electrical conductivity (ECse) of up to 5.3 dS/m,
and chickpea to 2.9 dS/m.
Conclusions As salinity increased, the effects of specif-
ic ion toxicity dominated, and water extraction by plants
Responsible Editor: W Richard Whalley.
A. R. Sheldon : R. C. Dalal : G. Kirchhof : P. M. Kopittke :
N. W. Menzies (*)
School of Agriculture and Food Sciences, The University of
Queensland, Qld, St Lucia 4072, Australia
e-mail: n.menzies@uq.edu.au
was significantly lower than that predicted on the basis
of the total soil water potential.
Keywords Soil water potential . Permanent wilting
point . Osmotic adjustment . Matric potential . Osmotic
potential
Introduction
Salinity is considered to reduce plant growth and per-
formance by a multitude of mechanisms including alter-
ations in water relations within the plant (Munns and
Tester 2008; Deinlein et al. 2014); ion deficiencies or
toxicities (Kingsbury and Epstein 1986; Ehret et al.
1990), and oxidative stress (Kravchik and Bernstein
2013; Shoresh et al. 2011; Reddy et al. 2015). Salinity
is also considered reduce the plant availability of soil
water, the osmotic effect on soil water potential, reduc-
ing plant water uptake (Clark 1990). Tolerance to salin-
ity may therefore involve variations in responses to each
of these factors (Munns and Tester 2008). This study
considers the impact of salinity on plant availability of
soil water. By decreasing the osmotic potential of the
soil solution (i.e. becoming more negative), plant access
to soil water is reduced because of the decrease in total
soil water potential. As the salinity of the soil increases,
so does the soil water content at which permanent
wilting point (PWP) is reached (Slayter 1957). From a
theoretical perspective, the PWP is determined by the
sum of matric and osmotic potentials. The effect of
salinity on soil water availability was modelled by
478
Groenevelt et al. (2004), based on the water character-
istic curve of the soil.
In order to maintain water uptake from a dry or saline
soil, plants must decrease their internal water potential
below that of the soil. This is done either by taking up
salts and compartmentalizing them within plant tissue,
or by synthesizing compatible organic solutes (Roy et al.
2014). The degree to which plants can adjust to dry and
saline conditions depends on their ability to maintain a
water potential below that of the soil (Reddy et al.
2015), and their ability to tolerate or exclude elevated
concentrations of NaCl present in the root environment
(Nemati et al. 2011). Current theory suggests that the
salt tolerance of the plant determines the degree to
which the plant is subject to the additional water stress
imposed by the osmotic component of soil water poten-
tial (Groenevelt et al. 2004). Plants with a high level of
salt tolerance will be able to overcome the additional
osmotic stress, and be subject to matric potential only;
like a tensiometer. Salt sensitive plants, however, will be
unable to overcome the osmotic potential, and are there-
fore subject to the total soil water potential (Groenevelt
et al. 2004).
Salinity not only affects the final soil water content
which plants can extract to, but also the rate at which
they use water (Cardon and Letey 1992; Homaee et al.
2002; Sepaskhah et al. 2006; Skaggs et al. 2006;
Fujimaki et al. 2008). Between field capacity and
PWP, two stages of water use are observed. At high
water contents, water use is constant until a threshold is
reached, from which the rate of water use decreases until
PWP is reached. Salinity reduces the rate of water use
during both stages (Homaee et al. 2002).
In the present study, a comparison was made between
two species, wheat (Triticum aestivum L.) and chickpea
(Cicer arietinum L.), in their ability to extract water
from a drying, saline soil. These species were chosen
as they are known to differ in their tolerance of salt. The
response of the species was then interpreted in the
context of the model presented by Groenevelt et al.
(2004). A range of other factors, such as Na and Cl
uptake are considered in order to examine the extent to
which potential growth reductions are due to the toxicity
of these ions, or additional water stress on the plant.
The soil used for this experiment was a Red Ferrosol
(Isbell 2002) (Oxisol, Soil Survey Staff 2010) topsoil
from Redland Bay in southeast Queensland, Australia.
This soil is a strongly structured heavy clay soil, acidic,
high in Fe-oxides and non-swelling. While a soil of this
Plant Soil (2017) 418:477–491
kind is not generally associated with salinity problems, it
was used for this study because adding NaCl would not
alter the physical behaviour of the soil. Therefore, the
effect of salt on plant water uptake would not be con-
founded with the effect of altered soil physical properties
on root proliferation and soil water potential. Further-
more, the relationship between soil water content and soil
solution composition for this soil has been characterized
(Dyer et al. 2008), affording a unique opportunity to
investigate the relative effects of water potential, plant
osmotic adjustment and specific ion toxicity.
Materials and methods
Soil water retention and modelling
The water retention curve for the soil was measured
using the pressure plate method (Klute 1986) to deter-
mine the relationship between water potential and gravi-
metric water content (GWC) for this soil, with eight
water contents ranging from saturation to PWP. Water
retention curves were determined at three salinity levels;
no added salt (control), control plus 1 g NaCl/kg, and
control plus 2 g NaCl/kg. The data for the control
treatment was used to model the water retention curve
for the soil using the equation described in Groenevelt
et al. (2004). Further modelling simulations were con-
ducted to examine the combined effect of matric poten-
tial and osmotic potential in this soil under saline con-
ditions (Groenevelt et al. 2004).
Pot trial
A pot trial was conducted in a glasshouse at The
University of Queensland (St Lucia, Australia) to ex-
amine the extent to which the effect of NaCl on PAW is
due to osmotic stress. The trial was arranged as two
(wheat and chickpea) randomized block design exper-
iments, consisting of a total of 60 experimental units,
with two plant species, 10 rates of NaCl, and three
replicates. Glasshouse maximum temperatures ranged
from 27 to 33 °C while heating was used to maintain
minimum temperature achieving a range from 15 to
21 °C. Relative humidity ranges from 52 to 74%.
Concurrent experiments were conducted with wheat
cv. Kennedy and chickpea cv. Jimbour grown in soil
contaminated with NaCl. Ten rates of NaCl were used,
from 0 to 7.5 g/kg (Table 1) – as later measured (see
Plant Soil (2017) 418:477–491 479

Table 1 Values of electrical conductivity of a saturation extract pots, and plant growth and foliar symptoms were mon-
(ECse), the osmotic potential at saturation, and the integral water
itored. Plants were harvested sequentially as each treat-
capacity for each of the NaCl treatments applied to a Red Ferrosol
soil ment reached PWP, as determined by early morning
foliar observations. If plants had not recovered turgor
NaCl ECse Osmotic potential Integral water overnight, they were destructively harvested that morn-
(g/kg) (dS/m) at saturation capacity* (g/g)
(−MPa) ing. For wheat, treatments up to ECse 1.66 dS/m were
harvested 9 d after watering ceased, treatments ECse
0.000 1.25 0.0452 0.271 2.06 to 2.87 dS/m after 11 d, treatments ECse 5.29 and
0.005 1.27 0.0458 0.271 10.9 dS/m after 13 d, and treatments ECse 17.4 and 25.4
0.125 1.66 0.0597 0.254 dS/m after 23 d, while for chickpea, treatments ECse
0.250 2.06 0.0742 0.240 1.25 and 1.27 dS/m were harvested 14 d after watering
0.375 2.46 0.0887 0.229 ceased, treatments ECse 1.66 to 2.46 dS/m after 15 d,
0.50 2.87 0.103 0.219 treatment ECse 2.87 after 19 d, and treatment ECse 5.29
1.25 5.29 0.191 0.180 after 29 d. Chickpea treatments ECse 10.9 to 25.4 dS/m
3.0 10.9 0.393 0.134 died. The harvested plant tops were dried to constant
5.0 17.4 0.626 0.106 weight at 65 °C, weighed, ground, then sub-samples
7.5 25.4 0.916 0.085 were digested in nitric-perchloric acid (Martinie and
*
Schilt 1976), and analysed by inductively coupled plas-
Calculated using Eq. 25, Groenevelt et al. 2004
ma atomic emission spectroscopy (ICPAES, Spectro
Ciros, Spectro Analytical Instruments, Kleve, Germany)
below), this yielded treatments with ECse values rang- for Al, B, Ca, Cu, Fe, K, Mg, Mn, Na, P, S and Zn. Plant
ing from 1.25 to 25.4 dS/m. samples were also analysed for Cl using nitric acid
Straight-sided pots (15 cm diameter, 20 cm deep) extraction and mercuric thiocyanate colorimetric analy-
were used in this experiment, lined with plastic bags. sis (Clesceri et al. 1998). Following harvest, the lower
Each pot had two layers of soil, the salt treatment and soil layer was placed in a plastic bag and roughly mixed.
half the basal nutrient application was applied to 3 kg of Moist soil sub-samples (≈100 g) were weighed, oven
soil in the lower layer, while the remaining basal nutrient dried and re-weighed for GWC, additional moist sam-
application was applied to the 500 g surface layer. Soil ples collected for analysis of water potential using a
was packed to 18 cm depth giving a bulk density of vapour-pressure psychrometer (Wescor HR-33 T with
1.1 g/cm3. In each case, the nutrients and salt were C-52 sample chamber, EliTech, Pureaux, France). The
added to enough water to bring the soil to field capacity, remaining soil was dried, mixed thoroughly and sub-
this solution was then added to the soil. Total basal sampled for electrical conductivity (EC) and pH mea-
nutrient application was (mg/pot) 1020 N, 180 P, surement on saturation extracts (ECse and pHse).
250 K, 105 Ca, 105 Mg, 18 Zn, 18 Cu, 190 Cl, 160 S, It is necessary to briefly define the terminology used
3.5 B, 0.8 Mo, as NH 4 NO 3 , KH 2 PO 4 , KCl, to describe EC. Unless otherwise stated, modelled
CaCl2.2H2O, MgSO4.6H2O, ZnSO4.7H2O, values for EC (termed ECs) are calculated for the con-
CuSO4.5H2O, H3BO3, and Na2MoO4.2H2O. Following dition when soil pores are saturated; Groenevelt et al.
salt and nutrient application, soils were allowed to equil- (2004) described this condition as corresponding rough-
ibrate for one week. ly to the conditions in a saturation paste extract. Exper-
Each pot was planted with five seeds, plants emerged imental values are for a saturation paste extract (termed
after four days in low salt treatments extending to 9 days ECse). The modelling assumes that all salts present
in the highest salt treatments; thinning to three uniform when soil pores are saturated remain in the soil solution
plants was undertaken three days after emergence. The as the soil dries (see detailed discussion of this assump-
soil surface was initially covered in a layer of 100 g of tion later); thus for a stated ECs condition, the modelled
white plastic beads to minimize evaporative loses, to soil solution EC will increase as the soil dries. Thus
which an additional 100 g was added at the final these two values, ECs (for modelled data) and ECse (for
watering, when plants were larger. Pots were maintained experimental data), are an indication of the total amount
at field capacity until 28 days after planting. At this time of salt in the soil, rather than the soil solution EC at each
watering ceased, water loss was monitored by weighing moisture content.
480
Statistical analysis
Statistical analysis of data in this study was conducted to
test for treatment effects, and identify differences in
plant responses. Analysis consisted of one-way
ANOVA with Tukey’s pairwise comparisons at a 95%
confidence interval (SAS/STAT ver 9.13., SAS Institute
Inc. 2004).
Results
Soil water retention and modelling
The soil water retention curve is shown in Fig. 1, to-
gether with line of best fit determined using eq. 6 from
Groenevelt et al. (2004). Addition of salt did not signif-
icantly alter the water retention curve (data not present-
ed). The curve fitting process optimises three parameters
to describe the fitted equation; n = 0.172, K0 = 1.67,
K1 = 0.826. An analysis of the soil using the approach of
Groenevelt et al. (2004) was undertaken (Fig. 2), with
soil water potential units in MPa, rather than m. Equa-
tions presented in Groenevelt et al. (2004), and used to
calculate data for Figs. 1 and 2 are listed in Table 2.
The function plotted in Fig. 2a indicates the ECs that
would be required to reach a total soil water potential
equivalent to PWP across a range of soil water contents.
For example, in a saturated soil, an ECs of approximate-
ly 42 dS/m would be required to create a total soil water
potential of −1.5 MPa. As the soil dries, the ECs value
represents the total amount of salt in the system, with the
osmotic potential contribution calculated assuming that
all of the salt present at saturation remains in the soil
Fig. 1 Water retention curve for the Red Ferrosol (Eq. 6,
Groenevelt et al. 2004). Points indicate treatment means with
standard deviations (n = 3)
Plant Soil (2017) 418:477–491
solution. Thus as the soil dries, the concentration of salt
in the soil solution increases, and the osmotic potential it
imposes is increasingly negative. Soil drying also makes
the matrix potential increasingly negative. As plants are
considered to respond to the total water potential, the
contribution of osmotic potential means that PWP is
reached at a lower matric potential as the total amount
of salt in the soil (ECs) is increased. Fig. 2b presents the
same relationship over a smaller range of ECs values
more relevant to plant growth; for any ECs the relation-
ship indicates the matrix potential which, combined
with the osmotic potential contributed by the salt present
in soil solution, will sum to PWP. The relationship
between EC s and soil matric potential shown in
Fig. 2a,b can then be transformed to show the relation-
ship between ECs and GWC (Fig. 2c), indicating that as
salt concentration increases the water content at PWP
also increases.
Differential water capacity represents the change in
GWC, with change in soil matric potential (Fig. 2dd),
and represents the capacity of the soil to hold or release
soil solution. For the Red Ferrosol, this indicates that at
high water contents, a relatively large change in soil
water content is associated with a small decrease in
matric potential, while a small decrease in soil water
content in a dry soil corresponds with a large decrease in
matric potential. No effect of EC or osmotic potential is
shown in Fig. 2d.
As the soil dries, the contribution of soil solution EC,
and the associated osmotic potential to total soil water
potential decreases (Fig. 2e). This occurs because soil
solution ionic strength (and the resultant osmotic pres-
sure) is negatively proportional to the soil solution vol-
ume, while the differential water capacity changes in
proportion to the logarithm of the soil solution volume.
Fig. 2e indicates the changing proportion of total soil
water potential due to osmotic potential as the soil dries
for a number of nominal ECs values. The region below
the line for each ECs value indicates the contribution of
matric potential, and the region above is osmotic poten-
tial. Increasing the salinity of the soil decreases the
availability of soil water to plants. This effect is greatest
at high water contents, as shown in Fig. 2f. Here the
combined effects of the physical properties of this soil
(Fig. 2d), where soil water is most readily available at
the lowest matric potentials (i.e., no water release point),
and the decreasing contribution of osmotic potential to
total water potential as the soil dries (Fig. 2e), are
shown. Fig. 2f also presents the total amount of water
Plant Soil (2017) 418:477–491
Fig. 2 Analysis of the Red Ferrosol using the approach of
Groenevelt et al. 2004. Graphs show, (a). the electrical conductiv-
ity of saturated soil, ECs, and the corresponding matric potential at
which PWP (−1.5 MPa) will occur (Eq. 12,), (b). the matric
potential at PWP (−1.5 MPa) for a relevant range of ECs
(Eq. 12), (c). the gravimetric water content at PWP (−1.5 MPa)
and the corresponding ECse (Eq. 14), (d). the differential water
available to plants, or integral water capacity, at each of
the ECs values shown, as the area under each curve
(quantified in Table 3).
In order to relate calculated values of GWC at PWP
to measured values, the GWC of the Red Ferrosol at
PWP was calculated for a range of NaCl concentrations
relevant to plant growth (Fig. 3c,d). The GWC at PWP
was calculated using three different methods. Firstly,
PWP was determined by the matric potential alone,
thereby ignoring any potential contribution from osmot-
ic potential (short-dash lines Fig. 3c,d). Secondly, the
PWP was determined by the sum of the matric potential
and the osmotic potential when it is assumed that the
ionic strength of the soil solution increases with decreas-
ing moisture content (solid lines Fig. 3c,d). Finally,
PWP was determined as the sum of the matric potential
481
capacity (Eq. 15), (e). the result of applying a weighting function
for the differential water capacity to account for osmotic effects on
soil water availability, accounting for the salt free condition, and
where ECs = 1, 2, 4, and 6 dS/m (Eq. 22), and (f). the variously
weighted differential water capacities, accounting for the salt free
condition, and where ECs = 1, 2, 4, and 6 dS/m (Eq. 24). Abbre-
viation: ECs, electrical conductivity at saturation
and the osmotic potential, with the osmotic potential of
the soil solution assumed to remain constant (i.e. not
decrease) as the soil dries (long-dash lines Fig. 3c,d).
This third scenario (long-dash line) most accurately rep-
resents the expected behaviour of the system based on
our previous observations of the effect of drying on the
soil solution composition of this soil, which indicated
that some salt adsorption was occurring in the range of
water contents between field capacity and −0.047 MPa
(Dyer et al. 2008). Salt adsorption occurs in variable
charge soils when an increase in soil solution ionic
strength increases the surface charge density, causing
an increase in adsorption of ions from the soil solution
onto the solid phase (Fotovat et al. 1997; Donn and
Menzies 2005; Dyer et al. 2008). In soils where salt
adsorption occurs due to variable charge minerals, or
482 Plant Soil (2017) 418:477–491

Table 2 Equations presented in Groenevelt et al. (2004), and used to calculate data for Figs. 1, 2, 3, 4, 5, 6, 7, 8

Equation


6 θðhm Þ ¼ θ150 þ k 1 ½expð−k 0 150Þ −exp −k 0 h−n
m 


12 EC s ðhm Þ ¼ ð150−hm Þð3:6θs Þ−1 ½θ150 þ k 1 ½expð−k 0 150−n Þ −exp −k 0 h−n
m 
 
14 1=n   −1=n
EC s ðθÞ ¼ θð3:6θs Þ−1 150−k 0 −ln expð−k 0 150−n Þ−k −1 1 ðθ−θ150 Þ

−ðnþ1Þ


15 C ðhmÞ≡− dhdθm ¼ nk 0 k 1 hm exp −k 0 h−n
m

 −2
 −1
22 ωomin ðhm ; hos Þ ¼ ½1 þ hos θs ðθ150 þ k 1 ½expð−k 0 150−n Þ −exp −k 0 h−n −ðnþ1Þ
m Þ nk 0 k 1 hm exp −k 0 h−n
m 
h
24
i−1
 −2 −1
C om ðhm ; hos Þ ¼ nk 0 k 1 h−mðnþ1Þ exp −k 0 h−n
m þ hos θs ðθ150 þ k 1 ½expð−k 0 150−n Þ −exp −k 0 h−n
m Þ 

∞
25 IWC ðhos Þ ¼ ∫0 C om ðhm ; hos Þdhm
Parameter Units Description
θ , θs g/g Soil water content: at saturation, at PWP (150 m)
hm , hos m Soil water potential: matric, and osmotic at saturation
ECs dS/m Soil EC at saturation
C 1/m Differential water capacity
ωomin (0–1) Weighting function
IWC g/g Integral water capacity

precipitation of sparingly soluble salts, such as gypsum, As the plants grew, those in the lowest NaCl treatments
the osmotic potential of the soil solution would not (ECse < 1.3 dS/m) showed the greatest effect of water
decrease linearly as the soil dries. restriction, while those in treatments from ECse 1.66 to
5.29 dS/m had increasingly severe foliar symptoms
associated with exposure to NaCl. Plants in mid-range
Pot trial NaCl treatments appeared to be less affected by wilting
than those in the lowest NaCl treatments which would
Foliar symptoms wilt in the middle part of the day, but then recover. The
plants in the treatments with ECse > 10 dS/m had much
Firstly, consideration is given to growth of wheat. Var- more severe foliar symptoms than those observed in the
iation in growth between the NaCl-treatments was evi- lower NaCl treatments. These foliar symptoms affected
dent from the time at which water stress was imposed, approximately half the plant in the ECse 10.9 dS/m
with plants grown at the highest three rates of NaCl treatment, and three quarters of the plant in the ECse
markedly smaller than those in the other treatments. 17.4 and 25.4 dS/m treatments. Growth was significant-
ly reduced in these treatments (Fig. 3a).
Table 3 Values of integral water capacity for ECse = 0, 1, 2, 4, and
6, for Red Ferrosol (Eq. 25, Groenevelt et al. 2004)
Next, consideration is given to the growth of chick-
pea. During the early part of the experiment, there was
ECse Osmotic potential at saturation Integral water capacity little visual difference in growth evident in treatments up
(dS/m) (−MPa) (g/g) to ECse 2.87 dS/m. Growth was substantially reduced in
0 0 0.678
plants grown in ECse 5.29 dS/m, and further reduced in
those grown in ECse 10.9 dS/m and greater (Fig. 3b).
1 0.036 0.286
Plants grown in the highest three NaCl treatments had
2 0.072 0.242
considerably impaired development, with small leaves
4 0.144 0.198
compared to those in lower NaCl treatments. These
6 0.216 0.171
plants developed necrosis of older leaves earlier than
ECse electrical conductivity of saturation extract other plants, and the end point of the experiment was
Plant Soil (2017) 418:477–491
Fig. 3 Plant dry weight; per pot for (a) wheat, and (b) chickpea;
values shown are treatment means with standard deviations
(n = 3). The gravimetric water content of soil at harvest for (c)
wheat, and (d) chickpea. Values shown are treatment means with
standard deviations (n = 3). Dashed line indicates PWP of matric
potential −1.2 MPa for wheat, and −0.9 MPa for chickpea (these
matric potential values were selected as they corresponded to the
final GWC in low-salinity treatments). Solid line indicates com-
bined matric and osmotic potential where all salts remain in the
determined by plant death rather than apparent wilting.
Across the range of NaCl treatments from ECse 1.66 to
5.29 dS/m, foliar symptoms associated with excess
NaCl were observed.
Water use
Plants were harvested when PWP was reached (or upon
plant death in highly saline treatments), as determined
by dawn observations. The GWC of the soil at harvest is
shown in Fig. 3c,d (data points). In the low-NaCl treat-
ments, the variation in PWP between wheat (ca. 0.17 g/
g, Fig. 3c) and chickpea (ca. 0.20 g/g, Fig. 3d) is due to
483
soil solution which increases in ionic strength. Long-dash line
indicates combined matric and osmotic potential where soil solu-
tion ionic strength remains constant as the soil dries, through
mechanisms of salt adsorption or precipitation. The soil water
potential at harvest as measured by vapour pressure psycrometer
(e) wheat, and (f) chickpea; values shown are treatment means
with standard deviations (n = 3). Abbreviation: ECse, electrical
conductivity of saturation extract
the difference in ability to extract water from the soil
under non-saline conditions.
Based upon the GWC, it was estimated that wheat
extracted water to a combined soil water potential of
−1.2 MPa, as determined by the total matric and osmotic
potential (i.e. solid line in Fig. 3c), at NaCl concentrations
up to 1.25 g/kg, corresponding to a soil ECse of 5.29 dS/
m. At NaCl additions > ECse 5.29 dS/m, wheat failed to
extract water to PWP (i.e. the measured gravimetric
moisture contents were higher than the solid line,
representing combined matric and osmotic potential,
Fig. 3c). The end point of the experiment for these plants
at elevated NaCl concentrations, as determined by either
wilting or death, was reached while considerable water
484
which in potential terms was considered available
remained in the soil. In the case of the ECse 25.4 dS/m
treatment, these plants were small and necrotic, and the
convergence of this point with the line representing PWP
may be due as much to passive wicking of water through
necrotic tissue as to transpiration losses.
Water extraction by chickpea was similar (Fig. 3d),
except that chickpea was able to extract water to PWP of
−0.8 MPa, as determined by combined matric and os-
motic potential, only up to NaCl concentrations of 0.5 g/
kg, or a soil ECse of 2.87 dS/m. Beyond this point, plants
apparently wilted or died before using the available
water, with measured GWC being substantially higher
than the combined matric and osmotic potential corre-
sponding to −0.8 MPa (solid line, Fig. 3d).
The total soil water potential, as measured by vapour
pressure psychrometer, is shown in Fig. 3e f. For both
wheat (Fig. 3e) and chickpea (Fig. 3f), the total soil
water potential at harvest was low (i.e. more negative)
at the lowest NaCl applications, but increased with
increasing ECse to a point of maximum soil water avail-
ability – corresponding to soil ECse values of 10.9 dS/m
for wheat and 5.29 dS/m for chickpea. Above these
points, water availability decreased (Fig. 3e,f), despite
the water content of the soil at harvest continuing to
increase (Fig. 3c,d). This point of inflection (Fig. 3e,f)
also coincides with the NaCl concentration at which
plants were unable to extract water to a soil water
content at harvest equivalent to PWP as determined by
combined matric and osmotic potential (Fig. 3c,d). At
the low range of NaCl additions, wheat extracted water
to soil water potentials in the range of −0.9 MPa to
−1.1 MPa (Fig. 3e), while chickpea extracted water to
total soil water potentials in the range of −0.6 to
−0.8 MPa (Fig. 3f). While some variability was ob-
served, there was no significant change in the soil water
potential to which wheat and chickpea extracted to, up
to ECse of 2.87 dS/m for wheat, and 2.06 dS/m for
chickpea. The values determined using a vapour pres-
sure psycrometer (Fig. 3e,f), which represent total
matric and osmotic potentials, are consistent with the
values calculated for the sum of matric and osmotic
potentials when all salts remained in soil solution
(Fig. 3c,d). As toxicity of NaCl increased, and plants
were unable to maintain water extraction, the availabil-
ity of the soil water remaining increased, to the peaks
observed at ECse of 10.9 dS/m for wheat, and 5.29 dS/m
for chickpea, beyond which, increasing NaCl decreased
water availability.
Plant Soil (2017) 418:477–491
The higher variability in water potentials observed in
the low NaCl treatments (Fig. 3)e,f, compared to water
contents (Fig. 3c,d) occurs because of the relationship
between soil water potential and soil water content for
this particular soil, where at low soil water content,
small changes in water content are associated with large
differences in water potential (Fig. 1). This effect is
observed again where daily water use is compared to
both soil water content and soil water potential (Fig. 4).
Daily water use by wheat (Fig. 4a,c,e), and chickpea
(Fig. 4bd,f) from the onset of water stress are shown in
relation to changing i) soil GWC, ii) soil matric poten-
tial, and iii) combined soil matric and osmotic potential.
Results are shown as two-segment, linear piecewise
regressions for daily water use for each treatment. Treat-
ments are grouped according to the pattern of response
shown to decreasing water availability. Results for treat-
ments with ECse > 10 dS/m have been omitted from
Fig. 4 for clarity. Initially a consistent relationship be-
tween soil ECse and plant water use was observed for
both wheat and chickpea, with plants in the lowest ECse
treatments having the highest water use. This is partic-
ularly evident in Fig. 4a,b. The relationship between soil
water availability and plant water use changed as ECse
increased. For wheat in NaCl treatments with ECse < 2.5
dS/m water use was initially high, and continued to
increase due to plant growth. Once soil water content
reduced to 0.3 g/g, rates of water use decreased rapidly.
Plants in ECse ranging from 2 to 5 dS/m had lower water
use initially (Fig. 4c,e), but as the soil dried were able to
maintain higher transpiration than plants grown in soil
with lower ECse. The ECse 5.29 dS/m treatment had
lower initial water use, due to osmotic stress, and re-
duced water use as the soil dried. For plants in treat-
ments with ECse ≥ 10.9 dS/m water use was low, and did
not vary with soil water content.
Water use by chickpea was initially related to the
amount of NaCl in the soil, with higher water use by
plants in lower NaCl treatments, as was observed for
wheat. Unlike wheat however, water use by chickpea
decreased slightly from the point at which watering
ceased (Fig. 4b), due to the comparatively slower
growth rate of chickpea. As the soil dried to water
contents less than 0.32 g/g, or matric potential <
−0.1 MPa, the variation in daily water use became
smaller, especially in treatments with ECse < 3 dS/m,
due to the increasing dominance of matric potential over
osmotic potential as the limiting factor affecting water
extraction. Chickpea plants grown in soil with an ECse
Plant Soil (2017) 418:477–491 485

Fig. 4 Daily water use of wheat

(a, c, e) and chickpea (b, d, f) in
relation to (a, b) GWC, (c ,d) soil
matric potential, and (e, f)
combined matric and osmotic
potential. Lines indicate two-
segment, linear piecewise
regressions for daily water use by
each treatment (n = 3). Note that
treatments with ECse > 10 dS/m
have been omitted for clarity.
Abbreviation: ECse, electrical
conductivity of saturation extract

of 5.29 dS/m had substantially lower water use than
those grown at lower NaCl concentrations, while plants
grown at ECse > 10 dS/m had low water use.
The transition between osmotic potential and matric
potential dominating rate of water use is evident when
comparing the rate water use relative to soil water con-
tent (Fig. 4b), with rate of water use relative to soil water
potential (Fig. 4d,f). Osmotic potential dominates rate of
water use at high soil water contents, with clear differ-
entiation between treatments according to soil EC
(Fig. 4b). As the soil dries, matric potential becomes
the dominant factor, and no clear differentiation between
treatments according to soil EC is evident (Fig. 4d,f).
Because of the relationship between the volume of soil
water, and the availability of the soil water (Fig. 2d), the
range of soil water contents over which soil water is
easily extracted by plants accounts for a small range in
soil water potential.
Plant growth
Dry weights were not reduced by NaCl treatments up to
ECse of 5.29 dS/m for wheat, and 2.87 dS/m for chick-
pea (Fig. 3a,b). Growth of wheat was reduced by 80% at
10.9 dS/m, and chickpea growth reduced by 50% at 5.29
dS/m. The growth response observed here is consistent
with the bent-stick model, long advocated by salinity
researchers, where little growth reduction occurred up to
a threshold salinity, above which a linear growth reduc-
tion occurred (Maas and Hoffman 1977). More recent
studies have shown the response of glycophytes to
salinity to be a linear decrease in growth with increasing
NaCl, without the initial plateau, and suggested that the
bent-stick response is an experimental artefact, rather
than a true physiological response (Deifel et al. 2006;
Kopittke et al. 2009). Indeed, we consider that in this
study the growth plateau that occurs at low EC is a type
486
of experimental artefact, and occurs due to the restricted
access to water by the plants. Those plants growing in
the lowest EC treatments had better growth initially, but
exhausted the water supply sooner while those at slight-
ly higher EC were able to maintain growth over a longer
period, so that no significant differences in growth were
observed over the low range of NaCl treatments.
Tissue Na and Cl concentrations
Tissue concentrations of Na showed no significant in-
crease with increasing soil NaCl, at soil NaCl concen-
trations up to 1.25 g/kg for wheat, and 0.5 g/kg for
chickpea, while plants in treatments with greater
amounts of NaCl had significantly higher tissue concen-
trations (P < 0.001) (Fig. 5). Tissue Cl concentrations in
wheat and chickpea increased with soil NaCl, but sig-
nificant differences from 0 NaCl were not observed until
soil concentrations were 1.25 g/kg for wheat and 0.25 g/
kg for chickpea. Tissue concentrations for plant nutri-
ents were not markedly affected by the treatments, and
were consistently above reported critical values (data
not provided).
Discussion
Plant water extraction
The response of wheat and chickpea to combined salt
and water stress indicates that the effect of these two
stresses on plant growth is both inter-related and com-
plex. The previous theory on the effect of osmotic stress
on soil water availability to glycophytic plants sug-
gested that, depending on the degree of salt tolerance
of the plant, osmotic adjustment would take place to
varying degrees (Groenevelt et al. 2004). The PWPs, as
determined by matric potential alone, and combined
matric and osmotic potentials, would mark the two
boundaries of maximum water extraction, creating a
wedge-shaped zone increasing in width with increasing
soil NaCl (for example, see Fig. 3c,d). A salt tolerant
plant would have a high level of osmotic adjustment and
extract water to a point close to PWP as determined by
matric potential (short-dash line in Fig. 3c,d), while a
salt sensitive plant would be unable to osmotically ad-
just, and therefore be able to extract less water, to a point
closer to PWP as determined by combined matric and
Plant Soil (2017) 418:477–491
osmotic potential (solid line in Fig. 3c,d) (Groenevelt
et al. 2004).
The growth response of plants to salinity has been
described in a two phase model, with Phase I being the
initial growth reduction caused by osmotic stress, and
Phase II occurring as Na and Cl accumulate in the
tissues, causing toxicity to the plant (Munns et al.
1995; Munns and Tester 2008). The results of this study
indicate that soil NaCl concentrations associated with
the onset of substantial accumulation of Na in the plant
tissues are also associated with a decrease in growth,
and the inability of plants to extract water to PWP. The
two phase model proposes that the onset of Phase II is
associated with the degree of salt tolerance of the plant,
and that more salt tolerant plants would encounter Phase
II at higher salt concentrations, or later in the growth
period (Munns et al. 1995; Munns and Tester 2008).
Wheat grown in soil with ECse > 5.29 dS/m had
decreased growth (Fig. 3a), increased tissue Na concen-
trations (Fig. 5a), and decreased water extraction
(Fig. 3c). Relatively more salt-sensitive chickpea expe-
rienced the same effects at ECse > 2.87 dS/m (Figs. 3b,d
and 5b). The relationship between plant growth, tissue
ion concentrations, and water extraction indicate that the
point at which plants are unable to extract water to PWP
may be an additional indicator of the onset of the phase
II. At low to moderate salinity, the maximum water
extraction that can be achieved by glycophytes is deter-
mined by the combined matric and osmotic potentials.
Variation in salinity tolerance is related to the range of
salinity over which extraction to PWP can take place. At
higher salinities, toxicity of NaCl becomes the dominant
factor, and plant growth, and water use are reduced
beyond the range predicted by the osmotic potential of
the soil solution.
Plant growth and water use
Many of these previous studies have been conducted in
nutrient solution, which although useful in understand-
ing plant physiological responses, exclude interactions
with the soil solid phase. For example, whilst nutrient
solution culture allows examination of the impact of
osmotic potential, it is not possible to investigate the
interaction between the plants and the soil (for example,
the interaction between osmotic and matric potentials),
as occurs in the field. This was the focus of the present
study.
Plant Soil (2017) 418:477–491
Fig. 5 The Na concentration in
plant tissue of (a) wheat, and (b)
chickpea, and the Cl
concentration in plant tissue of (c)
wheat, and (d) chickpea. Values
shown are treatment means with
standard deviations (n = 3).
Abbreviation: ECse, electrical
conductivity of saturation extract
Firstly, we give consideration to plants grown in
treatments where the final shoot dry mass was not
decreased by salinity (i.e. ≤ 5.29 dS/m for wheat, and
2.87 dS/m for chickpea, Fig. 3). These plants achieved
maximum water use, high growth, and low Na and Cl
tissue concentrations (Figs. 3, 4, and 5). For these plants,
although the increase in ECse was comparatively small
and did not decrease overall dry mass (Fig. 3a,b), the
salts did, however, reduce the rate of growth (data not
presented), and contributed to the reduced daily water
use (Fig. 4a,b) and the greater length of time required
before the onset of permanent wilting and harvest (for
wheat 9 d for low salt treatments c.f. 23 d for the highest
salt treatments, and for chickpea 14 d for low salt
treatments c.f. 29 d for the highest non-lethal salt treat-
ment). Across these low-salinity treatments, the GWC
measured at harvest remained relatively constant despite
increasing ECse (Fig. 3c,d) which was in contrast to the
slightly increasing PWP (when calculated as matric plus
osmotic potential, solid line in Fig. 3c,d). This data
suggests that plants were utilizing osmotic adjustment
to overcome the deleterious effects of decreasing osmot-
ic potential (PWP). Regardless, the magnitude of this
adjustment in these low-salinity treatments would be
small compared to that required at higher EC, and
whether or not osmotic adjustment overcame the dele-
terious decrease in osmotic potential is not clear, given
that the magnitude of the decrease in osmotic potential is
small relative to the experimental error (Fig. 3c,d). Fi-
nally, given that the relative contribution of the osmotic
487
potential to the total soil potential decreases as the soil
dries (Fig. 2e), the magnitude of the adverse effect of
salts (in terms of osmotic potential) in these low-salinity
treatments decreased even further as the soil dried
(Fig. 4a,b). Thus, for the plants in these non-toxic,
low-salinity treatments, (i) overall shoot dry mass was
not reduced, (ii) tissue concentrations of Na and Cl
remained low, (iii) plants achieved near-maximum water
use, but (iv) the rate of growth was reduced somewhat,
and (v) although the magnitude of any required osmotic
adjustment was small, it is likely to have contributed to
improved growth.
Next, we give consideration to treatments where
toxicity was first observed, being ECse 10.9 dS/m for
wheat and 5.29 dS/m for chickpea. For wheat, for ex-
ample, growth was decreased at ECse 10.9 dS/m by ca.
80%, compared to ECse 5.29 dS/m where there was
almost no reduction in growth (Fig. 3a). So, what caused
this marked reduction in growth for plants at ECse 10.9
dS/m? The osmotic potential decreased with increasing
salinity, but it was calculated to decrease PAWC from
0.24 g/g at an ECse of 5.29 dS/m (and 0.27 g/g at 0 dS/
m) to only 0.21 g/g at ECse 10.9 dS/m (Fig. 3c) – this
further 13% reduction in PAWC (caused by decreasing
osmotic potential) could not account for the ca. 80%
reduction in shoot dry mass. Indeed, the measured value
for GWC at ECse 10.9 dS/m (0.29 g/g, data-point,
Fig. 3c) was substantially higher than that calculated
for PWP, regardless of whether PWP was calculated
using both matric plus osmotic potentials (0.23 g/g,
488
solid line Fig. 3c) or only the matric potential (0.17 g/g
short-dash line Fig. 3c). In other words, the plants used
markedly less water than was available, even in the
complete absence of any internal osmotic adjustment.
This observation emphasises that the reduction in plant
growth was primarily due to a direct toxicity of NaCl
rather than due to osmotic stress. Indeed, it was noted
that the ECse values of 10.9 dS/m for wheat and 5.29 dS/
m for chickpea corresponded not only with substantial
reductions in growth, but also with marked increases in
tissue concentrations of Na and Cl (Fig. 5). Thus, for
these toxic treatments, (i) overall shoot dry mass was
reduced markedly, (ii) tissue concentrations of Na and
Cl increased sharply, (iii) water use was substantially
lower than what was available, and hence (iv) growth
was largely reduced due a direct toxicity of NaCl.
The findings of the present soil-based study extend
those of previous studies based in nutrient solutions. For
example, Munns and Tester (2008) state that Bthe os-
motic stress not only has an immediate effect on growth,
but also has a greater effect on growth rates than the
ionic stress. Ionic stress impacts on growth much later,
and with less effect than the osmotic stress, especially at
low to moderate salinity levels. Only at high salinity
levels, or in sensitive species that lack the ability to
control Na+ transport, does the ionic effect dominate
the osmotic effect.^ However, this is somewhat in con-
trast to the present soil-based study where water avail-
ability was influenced by the total water potential. Spe-
cifically, we found that at low salinities, decreases in
osmotic potential reduced the rate of growth, but not the
final dry mass production. Furthermore, at moderate
salinity levels, the reduction in growth was dominated
by ionic stress rather than osmotic stress, regardless of
whether osmotic adjustment was assumed to have oc-
curred or not.
The observations of the present study regarding ionic
stress (i.e. toxicity of NaCl) are in agreement with other
studies which highlight Na toxicity in wheat (Munns
et al. 1995; Munns 2002; Nuttall et al. 2006; El-
Hendawy et al. 2007), and chickpea (Lauter and
Munns 1986; Dua and Sharma 1997). Tissue Cl con-
centrations, and total Cl uptake increased steadily with
soil Cl concentrations, but were poorly correlated with
changes in growth and water use. Chloride is known to
be an important ion for osmotic adjustment in the plant,
particularly for stomatal function and maintaining turgor
pressure (Campbell 1996; Xu et al. 2000). While these
results suggest that Na may have a greater toxic effect on
Plant Soil (2017) 418:477–491
plant growth and function than Cl, because this experi-
ment uses NaCl confounding the effects of Na and Cl, it
is impossible to determine this conclusively.
The pattern of water use by wheat plants in treat-
ments of ECse 2 to 5 dS/m, where water use was initially
low but rates of water extraction were maintained for a
longer period than plants at lower and higher salinities
(Fig. 4), is attributed to the plants’ capacity to adapt to
stressful conditions. Osmotic adjustment has been dem-
onstrated in both wheat (Fischer et al. 2005) and in
chickpea (Morgan et al. 1991), and in this experiment
permitted plants grown at up to 5 dS/m salinity to extract
as much water as control plants. One concern with this
experiment is that the relatively rapid onset of water
stress that occurs in a pot-based system may have lim-
ited the extent to which osmotic adjustment could occur.
The impact of rate of onset of water stress has been
demonstrated for water extraction by Rhodes grass as a
result of the physical properties of the soil (Wehr et al.
2005). Plants grown in soils with low unsaturated hy-
draulic conductivity encountered intermittent water
stress earlier than plants grown in soils with high unsat-
urated hydraulic conductivity. Those plants with gradual
adaption to stress conditions, through intermittent water
stress, had longer survival and extracted to lower soil
water contents than plants grown under less stressful
conditions, which died quickly when water stress was
encountered (Wehr et al. 2005).
The results of this experiment, that the osmotic po-
tential caused by salinity dominates total water poten-
tial, and hence transpiration rates, at high soil water
contents, but as the plants use water the contribution of
the matrix potential rapidly exceeds the osmotic contri-
bution, correlates well with published results (Homaee
et al. 2002). Even when matrix potential dominates soil
water potential, published models suggest that the sep-
aration between treatments according to soil EC should
remain; i.e. that the importance of osmotic potential to
plant water use does not change as the soil dries. In the
case of this study, variation in transpiration rates accord-
ing to soil EC was not observed once transpiration rates
started to decrease due to water stress. This observation
may be related to a number of factors. Firstly, any small
variation in transpiration rate with salinity treatments
may be obscured by variation in environmental condi-
tions encountered in the glasshouse. Secondly, conver-
gence of transpiration rates would be expected as the
soil dries, due to the decreasing contribution of osmotic
potential to total soil water potential (Fig. 2e)
Plant Soil (2017) 418:477–491
(Groenevelt et al. 2004), a factor which is not adequately
considered in most transpiration models (Homaee et al.
2002; Fujimaki et al. 2008).
Salt adsorption and precipitation
At low salinities (ECse ≤ 5.29 dS/m for wheat, and
ECse ≤ 2.87 dS/m for chickpea), it appears that both
wheat and chickpea extracted water to the PWP estimat-
ed as the combined matric and osmotic potentials, where
the osmotic potential was assumed to be the concentrat-
ed soil solution, without adsorption or precipitation
reactions (Fig. 3). This observation, however, is not
statistically significant due to the small difference in
total soil water potential between a scenario with
100% concentration of soil solution (Fig. 3c,d: solid
line), and 100% adsorption/precipitation of soil solution
(Fig. 3c,d: long-dash line) at low salinities, and the
variability in plant water extraction. This indicates that
while reactions causing adsorption or precipitation in a
drying soil may significantly alter the contribution of
osmotic potential to total soil water potential in highly
saline soils, the magnitude of these effects are small over
the range of salinities where wheat and chickpea are able
to extract water to PWP.
The model proposed by Groenevelt et al. (2004) to
describe the effect of osmotic potential, caused by sa-
linity, on availability of soil water to plants, assumes that
salts present in the soil solution will be concentrated as
the soil dries, decreasing the osmotic potential of the soil
solution. Soils however, contain a variety of mineral and
organic surfaces on which exchange, precipitation and
adsorption reactions may occur as the soil dries. Thus
the soil solution may not become more concentrated in
direct proportion to the change in soil water content.
Methods for characterising the exact nature of the soil
solution in drying soils are limited and difficult, with
current techniques capable of extracting useful volumes
of soil solution to water potentials of −0.23 MPa, de-
pending on soil type (Dyer et al. 2008). Given that most
plants are capable of extracting soil water to −1.5 MPa,
there is a wide range of soil water potentials where the
exact nature of the soil solution composition is un-
known, and can only be estimated on the basis of
theoretical predictions using the soil chemical and phys-
ical characteristics. Nevertheless, data from existing
approaches show a range of soil solution concentration
changes with soil drying, with a number of processes
identified which result in the removal of salt from the
489
soil solution. These include salt adsorption and precip-
itation reactions.
Salt adsorption occurs due to the interaction between
ions in the soil solution and variable charge minerals in
the soil. As the ionic strength of the soil solution in-
creases, the surface charge density on variable charge
mineral surfaces increases. This increase in surface
charge density results in an increase in uptake of ions
from the solution, and adsorption onto the solid phase. A
soil with a combination of positively charged sites on Al
and Fe oxides, and negatively charged sites on clay
minerals and organic matter, has the potential to increase
in both cation and anion exchange capacity with increas-
ing ionic strength. When increases in cation and anion
exchange capacity of equivalent magnitude occur, cat-
ions and anions of the soil solution are adsorbed onto the
solid phase in equal amounts (Qafoku and Sumner
2002; Dyer et al. 2008). Salt adsorption and precipita-
tion have a similar net effect on the concentration of the
soil solution as the soil dries. Precipitation of sparingly
soluble salts in drying soils can result in a change in soil
solution composition, and a smaller increase in solution
concentration than would be predicted from the simple
concentration-dilution model. A range of sparingly sol-
uble salts, particularly carbonates, bicarbonates and sul-
phates, are commonly present in soils. Gypsum is of
particular interest due to its common occurrence in
saline soils, the relatively large quantities it occurs in,
and the degree of solubility which is sufficient to create
the potential for significant measurement errors (DNRQ
1997).
Conclusions
1. At low salinities, although growth rate was reduced,
the maximum water use was not. This appeared to
occur because of osmotic adjustment, with PWP
determined by the sum of matric and osmotic
potentials.
2. At these low salinities, deleterious osmotic effects
were important in reducing growth rate at high
moisture contents, but as soils became drier, the
relative contribution of osmotic effects decreased
and water availability was dictated largely by matric
potential.
3. At moderate salinities that reduced plant biomass
production, osmotic effects appeared to be small,
with the main reduction in biomass due to ionic
490
stress (NaCl). This was evidenced by the observa-
tion that considerable available-water remained
within the soil (irrespective of the method used to
calculate PWP).
4. The point at which plants are unable to maintain
water extraction to PWP as determined by the sum
of matric and osmotic potentials is proposed as an
additional indicator of the onset of the toxic phase of
salinity.
From a pragmatic standpoint these finding can be
used to guide the way that we go about measuring and
interpreting soil water potential. There are various ways
in which we are readily able to determine soil water
potential directly (tensiometer, gypsum blocks), or indi-
rectly through the use of routine measurement of soil
water content (TDR, gravimetric moisture content) in
combination with an established relationship between
soil water content and soil matric potential (tension
tables / pressure plate). In contrast, total water potential,
or the osmotic potential, is difficult to measure. The
results of this study indicate that, for this soil at least,
measurement of matric potential alone is sufficient. At
low soil salt contents, osmotic potential is small relative
to matric potential throughout the range of plant avail-
able water contents, therefore a measurement of matric
potential alone is sufficient. As the soil salt content
increases, specific ion toxicity limits plant growth before
osmotic potential makes a substantial contribution to
soil water potential. In this situation, measurements
indicating the soil salt content, such as EC or extractable
Na or Cl, will be more effective indicators of plant
performance than measures of soil water potential.
References
Campbell NA (1996) Biology. Benjamin/Cummings Publishing
Co. Inc., Menlo Park
Cardon GE, Letey J (1992) Plant water-uptake terms evaluated for
soil-water and solute movement models. Soil Sci Soc Am J
56:1876–1880
Clark GA (1990) Measurement of soil water potential. HortSci 25:
1548–1551
Clesceri LS, Greenberg AE, Eaton AD (1998) Standard methods
for the examination of water and wastewater. United Book
Press Inc., Baltimore
Deifel KS, Kopittke PM, Menzies NW (2006) Growth response of
various perennial grasses to increasing salinity. J Plant Nutr
29:1573–1584
Plant Soil (2017) 418:477–491
Deinlein U, Stephan AB, Horie T, Luo W, Xu G, Schroeder JI
(2014) Plant salt-tolerance mechanisms. Trends in Plant Sci
19:371–379
DNRQ (1997) Salinity management handbook. Queensland
Department of Natural Resources. Scientific Publishing,
Brisbane
Donn MJ, Menzies NW (2005) The effect of ionic strength vari-
ation and anion competition on the development of nitrate
accumulations in variable charge subsoils. Aust J Soil Res
43(1):43
Dua RP, Sharma SK (1997) Suitable genotypes of gram (Cicer
arietinum) and mechanism of their tolerance to salinity.
Indian J Agr Sci 67:440–443
Dyer CL, Kopittke PM, Sheldon AR, Menzies NW (2008)
Influence of soil moisture content on soil solution composi-
tion. Soil Sci Soc Am J 72:355–361
Ehret DL, Redmann RE, Harvey BL, Cipywnyk A (1990)
Salinity-induced calcium deficiency in wheat and barley.
Plant Soil 128:143–151
El-Hendawy SE, Hu Y, Schmidhalter U (2007) Assessing the
suitability of various physiological traits to screen wheat
genotypes for salt tolerance. J Integr Plant Biol 49:1352–
1360
Fischer RA, Sayre KD, Reynolds MP (2005) Osmotic adjustments
in wheat in relation to grain yield under water deficit envi-
ronments. Agron J 97:1062–1071
Fotovat A, Naiduavendra, Sumner ME (1997) Water : soil ratio
influences aqueous phase chemistry of indigenous copper
and zinc in soils. Aust J Soil Res 35(4):687
Fujimaki H, Ando Y, Cui YB, Inoue M (2008) Parameter estima-
tion of a root water uptake model under salinity stress.
Vadose Zone J 7:31–38
Groenevelt PH, Grant CD, Murray RS (2004) On water availabil-
ity in saline soils. Aust J Soil Res 42:833–840
Homaee M, Feddes RA, Dirksen C (2002) A macroscopic water
extraction model for non-uniform transient salinity and water
stress. Soil Sci Soc Am J 66:1764–1772
Isbell RF (2002) The Australian soil classification. CSIRO
Publishing, Melbourne
Kingsbury RW, Epstein E (1986) Salt sensitivity in wheat. A case
for specific ion toxicity. Plant Physiol 80:651–654
Klute A (1986) Water retention: laboratory methods. In: Klute a,
methods of soil analysis. Part 1. Physical and mineralogical
methods, 2nd edn., American Society of Agronomy / soil
science Society of America, Madison, pp. 635-662
Kopittke PM, Kopittke RA, Menzies NW (2009) Measurement
and interpretation of salinity tolerance in four perennial
grasses. J Plant Nutr 32:30–43
Kravchik M, Bernstein N (2013) Effects of salinity on the tran-
scriptome of growing maize leaf cells point at cell-age spec-
ificity in the involvement of the antioxidative response in cell
growth restriction. BMC Genomics 14:24
Lauter DJ, Munns DN (1986) Salt resistance of chickpea geno-
types in solutions salinized with NaCl or Na2SO4. Plant Soil
95:271–279
Maas EV, Hoffman GJ (1977) Crop salt tolerance - current assess-
ment. J Irr Drain Div -ASCE:115–131
Martinie GD, Schilt AA (1976) Investigation of the wet oxidation
efficiencies of perchloric acid mixtures. Anal Chem 48:70–
74
Plant Soil (2017) 418:477–491
Morgan JM, Rodreiguez-Maribona B, Knights EJ (1991)
Adaption to water-deficit in chickpea breeding lines by os-
moregulation: relationship to grain-yields in the field. Field
Crops Res 27:61–70
Munns R (2002) Comparative physiology of salt and water stress.
Plant Cell Environ 25:239–250
Munns R, Schachtman DP, Condon AG (1995) The significance
of a two-phase growth response to salinity in wheat and
barley. Aust J Plant Physiol 22:561–569
Munns R, Tester M (2008) Mechanisms of salinity tolerance. Ann
Rev Plant Biol 59:651–681
Nemati I, Moradi F, Gholizadeh S, Esmaeili MA, Bihamta MR
(2011) The effect of salinity stress on ions and soluble sugars
distribution in leaves, leaf sheaths and roots of rice ( Oryza
sativa L.) seedlings. Plant Soil Environ 57:26–33
Nuttall JG, Armstrong RD, Connor DJ (2006) Early growth of
wheat is more sensitive to salinity than boron at levels en-
countered in alkaline soils of south-eastern Australia. Aust J
Exp Ag 46:1507–1514
Qafoku NP, Sumner ME (2002) Adsorption and desorption of
indifferent ions in variable charge subsoils: the possible effect
of particle interactions on the counter-ion charge density. Soil
Sci Soc Am J 66:1231–1239
Reddy PS, Jogeswar G, Rasineni GK, Maheswari M, Reddy AR,
Varshney RK, Kishor PBK (2015) Proline over-
accumulation alleviates salt stress and protects photosynthet-
ic and antioxidant enzyme activities in transgenic sorghum
491
(Sorghum bicolor (L.) Moench). Plant Physiol Biochem 94:
104–113
Roy SJ, Negrao S, Tester M (2014) Salt resistant crop plants. Curr
Opin Biotech 26:115–124
SAS Institute Inc. (2004) SAS/STAT Version 9.13. SAS Institute
Inc., Cary, North Carolina
Sepaskhah AR, Bazrafshan-Jahromi AR, Shirmohammadi-
Aliakbarkhani Z (2006) Development and evaluation of a
model for yield production of wheat, maize and sugarbeet
under water and salt stresses. Biosyst Eng 93:139–152
Shoresh M, Spivak M, Bernstein N (2011) Involvement of
calcium-mediated effects on ROS metabolism in the regula-
tion of growth improvement under salinity. Free Radic Biol
Med 51:1221–1234
Skaggs TH, Shouse PJ, Poss JA (2006) Irrigating forage crops
with saline waters: 2. Modelling root uptake and drainage.
Vadose Zone J 5:824–837
Slayter RO (1957) The significance of the permanent wilting
percentage in studies of plant soil water relations. Bot Rev
10:585–636
Soil Survey Staff (2010) Keys to soil taxonomy 11th edn. USDA -
Natural Resources Conservation Service, Washington, DC
Wehr JB, So HB, Menzies NW, Fulton I (2005) Hydraulic prop-
erties of layered soils influence survival of Rhodes grass
(Chloris gayana Kunth.) during water stress. Plant Soil
270:287–297
Xu G, Magen H, Tarchitzky J, Kafkafi U (2000) Advances in
chloride nutrition of plants. Adv Agron 68:97–150