South African Journal of Plant and Soil

ISSN: 0257-1862 (Print) 2167-034X (Online) Journal homepage: http://www.tandfonline.com/loi/tjps20

Effects of sodium sulphate, sodium chloride and

manganese sulphate on kikuyu (Pennisetum
clandestinum) growth and ion uptake

A. J. Mills , M. V. Fey & C. E. Johnson

To cite this article: A. J. Mills , M. V. Fey & C. E. Johnson (2004) Effects of sodium sulphate,
sodium chloride and manganese sulphate on kikuyu (Pennisetum clandestinum)
growth and ion uptake, South African Journal of Plant and Soil, 21:4, 209-213, DOI:
10.1080/02571862.2004.10635051

To link to this article: http://dx.doi.org/10.1080/02571862.2004.10635051

Published online: 15 Jan 2013.

Submit your article to this journal

Article views: 42

View related articles

Full Terms & Conditions of access and use can be found at

http://www.tandfonline.com/action/journalInformation?journalCode=tjps20

Download by: [92.96.139.115] Date: 31 January 2016, At: 01:11

 S. Afr. J. Plant Soil 2004, 21 (4) 209

Effects of sodium sulphate, sodium chloride and manganese sulphate

on kikuyu (Pennisetum clandestinum) growth and ion uptake

'II 1 2* , M .V . Fey'
A .J. MIS' " J 0 hnson 3
1 an deE
1 Department of Soil Science, University of Stellenbosch, Private Bag X01,
Stellenbosch, 7602, South Africa .
2National Botanical Institute, Private Bag X7, Claremont, 7735, South Africa
3Department of Civil and Environmental Engineering, Syracuse University,
Syracuse, New York, 13244, USA

Accepted 24 February 2004

Irrigation with saline water is increasingly practised yet an understanding of plant response to different salt types
remains largely elusive. Our study investigated the effects of irrigation with simulated effluent containing three
salts on the growth and composition of a common South African pasture grass. We treated pots of kikuyu (Pen-
nisetum clandestinum Hochst. ex Chiov.) in a controlled environment with solutions of sodium sulphate
(Na2S04) or chloride (NaCI) at saturated soil solution electrical conductivity (EC) of 0,5, 10 and 20 dS m- 1 , com-
bined with 0, 900 or 1800 mg kg- 1 Mn as MnS04' Kikuyu yields in the Na2S04 treatments were significantly
lower than in corresponding NaCI treatments. This could be explained by increased ion imbalance and osmotic
stress as a result of higher ionic strength and a greater concentration of neutral ion pairs in the Na2S04 system
at similar EC. An apparent Na2S04-induced Ca deficiency was attributed to suppressed Ca 2+ activity through
Downloaded by [92.96.139.115] at 01:11 31 January 2016

ion pairing at high ionic strengths. Under the conditions of this experiment ionic strength appeared to be more
effective than EC as a measure of salinity stress when comparing the effects of S04 and Cion growth of kikuyu.

Key words: ionic strength, Pennisetum clandestinum, salinity, sodium chloride, sodium sulphate

* To whom correspondence should be addressed (E-mail: mills@nbi.ac.za)

Introduction
Irrigation with saline waters is becoming increasingly com-
mon as agriculture expands into marginal areas with lower
quality water (Ayers & Westcot, 1985; Awada et al., 1995).
Furthermore, because of stringent water quality regulations, it
is difficult and expensive to discharge saline water to surface
water, making irrigation an attractive management alternative
(Devitt,1989).
A thorough understanding of salinity effects on plants and
soils remains largely elusive (Cheeseman, 1988) and is con-
founded by the highly variable nature of salinity effects on
plant growth (Curtin, Steppuhn & Selles, 1993). The relative
sensitivity of plants to Na2S04 versus NaCI salinity, for
example, varies considerably according to the plant species
under investigation (Mengel & Kirkby, 1978; Paek, Chandler
& Thorpe, 1988; Manchanda & Sharma, 1989; Warne et aI.,
1990; Awada et al., 1995). The explanations proposed for the
varied responses include physiological differences between
plant species or even cultivars (Datta et al., 1995), as well as
ionic differences between salt types and the consequent
effects on nutrient uptake (Sposito, 1989). Furthermore, the
time frame of response measurement can be important in
determining the dominant response mechanism which can be
dual, with osmotic or water stress being followed by internal
injury (Munns, 2002). ' I
This study was prompted by the proposed disposal of'
three saline effluents, dominated by Na 2S04, NaC!, or MnS04,
onto kikuyu (Pennisetum clandestinum Hochst. ex Chiov.)
pasture in Mpumalanga Province, South Africa. Kikuyu is
known to be highly tolerant of both NaCI (Russell, 1976) and
MnCI 2 (Rayment & Verrall, 1980). The response of this grass
to other salt types has not, however, been documented. Our
objectives were to compare the effects of Na2S04 and NaCI,
and their interaction with MnS04 on the growth and composi-
tion of kikuyu and to develop a better understanding of the
salinity response mechanism by studying the chemical com-
position of the soil solution.
Materials and methods
Kikuyu was grown from I dm 3 sods in non-draining pots in a
loamy soil (23% clay) containing 3.8% organic matter taken
from a road verge on the University of Cape Town campus.
Basal nutrients were added to the soil in the following quanti-
ties, per kg dry soil: 100 mg P as NaH 2P04.H20; 31 mg Mg
as MgS04.7H20; 49 mg K as KCI, and 200 mg N as
NH4N0 3. Salinity effects of Na2S04, NaCI and MnS04 were
determined in a factorial experiment involving 24 treatments
in a randomised block design with two replications. Soils
were treated before planting with sufficient Na2S04 or NaCI
to produce a range of electrical conductivity (EC) in the satu-
ration extract of the soil (corresponding to a water content of
430 cm3 kg- I dry soil) up to a maximum of 21 dS m- I . In
addition, three levels of MnS04 (0,900 and 1600 mg Mn kg- I
dry soil) were superimposed on each of the series of Na salt
treatments. The concentration and volume of salt solution
added to soils in each treatment are presented in Table I. The
grass was maintained on a daily regime of II hours artificial
lighting (sodium halide and incandescent) and temperature
control between 20°C (night) and 28°C (day). Pots were irri-
3
gated with tap water every day to a field capacity of 170 cm
I
kg- until harvesting after 3 weeks of growth. Shoot mass was
determined after oven drying at 65°C. Milled shoots were
ashed at 480°C and dissolved in 5 MHCl at 50°C for 30 min-
utes. The diluted digests were analysed by direct current
plasma (DCP) optical emission spectrometry for Ca, Mg, Na
and K. Saturated paste extracts (Rhoades, 1982) were pi'e;.
 210 S. Afr. Tydskr. Plant Grond 2004, 21 (2)

pared from 400 g dry soil taken from each pot of treated soil stress on plant growth than NaCI solutions at the same EC.
prior to planting. The extracts were analysed for EC, pH, Mn This can be mechanistically explained by ion speciation of
by atomic absorption spectrometry, and other soluble cations the sulphate-dominated soil solutions, which revealed that up
(Ca z+, MgZ+, K+, NH4+ and Na+) and anions (Cr, S042- and
to 57% of cation species are bound as neutral ion pairs (e.g.
NOT) by ion chromatography using CS 12 and Ionpac AS4A
CaS04 and MgS0 4; Table 3). These ion pairs contribute to
columns on a Dionex DX300 instrument. Data were analysed
using a multifactorial ANOV A design and the software pack- osmotic stress but do not affect EC which, as has been previ-
age Unistat 5.0. Salinity treatment effects were assessed ously suggested (Papadopoulos, 1986), consequently is
using the Tukey-HSD test. Equilibrium ionic strength, ion unsuitable as an index of osmotic stress in sulphate-domi-
speciation and the activity of free ions in solution were com- nated solutions.
puted using the PHREEQC ion speciation model (Parkhurst
& Appe10, 1999). Table 2 Mean kikuyu yields (g dry matter pot-1) under
different treatments of sodium sulphate, sodium chloride and
Table 1 Concentration and volume of Na2S04, NaCI manganese sulphate. Values are means of the two replicates
and MnS04 solutions added to soils of each treatment for each treatment
Solution Level Volume added Dry matter yield (g poe!)
Treatment (g 1- 1) (dS m- 1)
Manganese treatment Mean(!)
(ml kg-1 soil) Sodium sulphate(2) 0 800 1600
Na2S04 14.2 -5 125 o 2.7 3.6 2.6 3.0 a
5 2.4 2.6 2.8 2.6 ab
Downloaded by [92.96.139.115] at 01:11 31 January 2016

31.8 -10 125

to 1.9 2.0 1.1 I.7bc
33.6 -20 250 20 1.0 1.4 1.3 1.2 c
NaCI 32 -5 25 Mean 2.0 2.4 1.9 2.1
70 -10 25 Sodium chloride
o 3.1 3.4 3.0 3.2 a
147 -20 25
5 2.1 2.8 3.4 2.8 ab
(g Mn kg-1) 10 2.6 2.2 2.1 2.4 ab
20 1.9 1.9 1.8 1.9 b
MnS04· H20 549 0.9 5
Mean 2.4 2.6 2.6 2.5
1.8 10 ANOVA, F-Value
Salt type Sodium chloride
Results and discussion Sodium sulphate 31.5 (P<O.OOI)
(I)Means within a column followed by the same letters are not signif-
icantly different at P=0.05 (Tukey-HSD test); (2)Four salinity treat-
Yield patterns ments: no addition of salt; addition of salt to achieve an electrical
Kikuyu shoot yields in the NaZS04 treatments were signifi- conductivity of -5, -10, -20 dS m- I in a saturated paste extract.
cantly lower than in NaCl treatments (P<O.OO I), while Three manganese treatments: 0, 900 and 1600 mg Mn kg-I.
MnS04 had no significant effect in any of the treatments
(P>0.05) (Table 2). Yields decreased in a curvilinear manner
with an increase in EC (Figure 1). A good measure of salinity Table 3 Ca speciation in saturated paste extracts of selected
stress would reflect an equal increase in salinity stress in treatments, as calculated using PHREEQC
response to a given increment of all salt types. Electrical con-
ductivity, the conventional measure of salinity stress (Camp-
bell, Bower & Richards, 1948), does not meet this c;
requirement for our comparison ofNaz S04 and NaCl salini-
ties. The inadequacy of EC is demonstrated by the results in
U
~"7'-"
= se ~ ~
.-:'::
~ ~ ~ ~ ~
Figure 1 and might be explained by additional osmotic and/or ~! ~ ~!
ionic stresses imposed by the S04 ion. Sodium sulphate treatment (no manganese addition)
o 6.4 6 0.4 3.1
Osmotic stress: a function of total solute concentration 0.5 10.2 6.4 3.8 2.6
The normally linear relationship between EC and ionic 12.7 6.8 5.9 2.3
strength, as reported for different soil solutions by Snoeyink
2 14.5 6.2 8.3 1.8
& Jenkins (1980) appears to be a poor model for solutions
Sodium sulphate treatment (Plus 1600 mg Mn kg-I)
dominated by Na ZS04 or NaCl. The total solute concentration
(including uncharged ion pairs) of Na2S04 solutions in our 2 12.5 5 7.5 1.3
experiment was up to 40% greater than that ofNaCl solutions Sodium chloride treatment (no manganese addition)
at the same EC. Ionic strengths were up to 53% greater in o 6.6 6.2 0.4 3.3
Na ZS04 solutions than in NaCl solutions at the same EC.
0.5 10.4 10.1 0.3 4.4
Osmotic potential is directly proportional to total solute con-
12.4 12.1 0.3 4.6
centration (Kramer, 1983) and thus the higher solute concen-
tration of Na2S04 solutions should impose a greater osmotic 2 14.3 14 0.3 4.8
 S. Afr. 1. Plant Soil 2004, 21(2) 211

We thought that total solute concentration (a summation of Calcium and S04 interactions
major cations and anions in a saturated paste extract) might Calcium availability is of critical importance for plants cop-
better reflect osmotic potential than EC. When using this ing with salinity stress because it is a structural component of
measure, however, lower yields at the same solute concentra- the plasmalemma, playing a role in controlling cell membrane
tion in Na2S04 treatments compared with NaCI treatments permeability and selectivity (Maas & Grieve, 1987; Hansen
are still evident (Figure 2). This pattern could possibly be & Munns, 1988; Suhayda et ai., 1992; Gratton & Grieve,
explained by an additional salinity stress factor associated 1999). Sodium-induced Ca deficiency has been cited as a
with ion imbalance and the effect ofS0 4 on Ca uptake. major cause of reduced yield under Na salinity stress (Maas
& Grieve, 1987; Awada et ai., 1995; Ashraf, Arfan and
4.5 Ahmad, 2003; Kaya, Ak & Higgs, 2003). Kikuyu did not
4 exhibit a marked reduction in Ca uptake with increasing NaCI
..•.. Sodium sulphate
3.5 stress (Figure 3), which suggests that it is Na-tolerant in this
- 0 - Sodium chloride
'-
0 3 respect. Sodium sulphate salinity stress, however, resulted in
Q.

.9 2.5 ~ a highly significant reduction (P<O.O I) in Ca foliar level (Fig-

"0
Qj 2 ........ ure 3), which could therefore, as has been suggested previ-
'>'
~ 1.5 " .... -._-. ously (Janzen & Chang, 1987), be the main cause of lower
0
yields in Na2S04 treatments.
0.5 Reduced Ca uptake in Na2S04 treatments might be
0 .--- accounted for by higher ionic strength and a different compl-
5 7 9 11 13 15 17 19 21
Downloaded by [92.96.139.115] at 01:11 31 January 2016

exation chemistry of Ca. Ion activity in aqueous solution is

Electrical conductivity (dS m") reduced by increasing ionic strength and the reduction is
greater for ions of higher valence (Snoeyink & Jenkins,
Figure 1 Dry matter yield versus electrical conductivity of saturated
paste extracts in Na2S04 and NaCI treatments. Data presented are
1980). For example, Ca activity is reduced by more than 50%
from experiments conducted with 900 mg Mn added kg-I dry soil. as ionic strength approaches 0.1 M, compared with its activity
Error bars depict values from the two replicates in each treatment. in an infinitely dilute solution (Snoeyink & Jenkins, 1980).
Solutions of Na2S04 in our experiments resulted in ionic
4.5 strength exceeding 0.4 M at the highest salinity level. The for-
4 mation of neutral ion pairs (CaS04o) lowers Ca 2+ activity
"'"[ 3.5 still further in Na2S04 treatments (Papadopoulos, 1986; Jan-
'"E 3 zen & Chang, 1987). Ion speciation calculations from satu-
~ 2.5· rated paste extract data using the PHREEQC model suggested
.s
3;! 2 that 57% of Ca is bound in CaS040 ion pairs at the highest
.~
~ 1.5
1 ~
..... Na2S04 treatment (Table 3). By contrast, only 2% of Ca is
o
\ ...•... Sodium sulphate bound in CaS040 at the highest NaCI treatment. Free Ca z+
0.5 - -0- Sodium chloride activity was consequently much greater in the highest NaCI
o treatment than the highest NaZS04 treatment (4.8 vs 1.8;
0.08 0.18 0.28 0.38
Table 3). We therefore attribute reduced Ca uptake observed
Total solute (mol r')
at higher Na2S04 salinity levels (Figure 3) to reduced activity
Figure 2 Dry matter yield versus total solute concentration (summa- of free Ca2 + ions, while recognising that in soil drier than sat-
tion of the concentration of all major cations and anions in saturated uration other factors, such as cation exchange and gypsum
paste extracts) for Na2S04 and NaCI treatments. Data presented are precipitation, might also have played a role in limiting the
from experiments conducted with 900 mg Mn added kg-' dry soil. supply of Ca to plant roots. In this context, it is worth record-
Error bars depict values from the two replicates in each treatment. ing that the calculated saturation index for gypsum in the sul-
phate-dominated solutions was close to zero, suggesting the
0.33 probability of gypsum formation under drier conditions e.g. at

. ·' ": :
(NaCll)
field capacity. Nevertheless it is evident from Table 3 that

~
0.28
····_·······f (NaCI2)
total Ca in solution at saturation is effectively identical in
. --1 both salt systems.

tW
(N.CI3)
~
.!!!
0.23
1.. .. ···'-1. . ----T ~ Calcium uptake was also reduced by the application of
MnS04' This we attributed to an even greater reduction in
~ I " ' : ... ,."
Ca z+ activity, brought about by an even greater S04 concen-
.••. ••..• •.••.•.•..•..•.•.• (Na,SO,1)
0.18
.. ..::.:........... ··········f (NB,SO.2)
tration and hence CaS040 ion pairing. The lowest Caz+ activ-
······l···········.. · .......·...~.t:! ~_a,SO.3t '
ity was recorded in the treatment containing the most Na2S04
0.13 -
3 13 18 and MnS04 (Table 3).
Electrical conductivity (dS m")
S04 effects on osmotic adjustment
Figure 3 Foliar Ca versus electrical conductivity for Na2S04 and
NaCI treatments. Data from all Mn addition levels are included: I =
Increased tissue concentration of osmotically active solutes
a mg Mn kg-I; 2 = 900 mg Mn kg-I; 3 = 1600 mg Mn kg-I. There is a (osmotic adjustment) is frequently utilised by salt-tolerant
missing data point for NaCI (2) due to a laboratory error. Error bars plants to counteract the osmotic stress of salinity (Yeo, 1983).
depict values from the two replicates in each treatment. Total cation uptake by kikuyu increased markedly with an
 212 S. Afr. TYdskr. Plant Grond 2004,21(2)

increase in salinity in both NaCI and Na2S04 treatments, yet
cation uptake was significantly lower in Na2S04 treatments at
the same EC level (Figure 4) suggesting that osmotic adjust-
ment was less effective in the latter. One explanation for this
may lie in a reduced activity of cations as was the case with
Ca. Magnesium and K also showed reduced uptake in
Na2S04 treatments relative to NaCI treatments. An alterna-
tive explanation may be found in the effect of Cl. Chloride is
known as a highly effective ion for osmotic adjustment pur-
poses, particularly in salt-tolerant grasses (Peacock, Dudeck
& Wildmon, 1993). It is thus possible that kikuyu uses CI for
osmotic adjustment and increased uptake of CI in the high
NaCI treatments consequently facilitated foliar cation enrich-
ment.
1.91
~; 1.8
J
§. 1.7
2 1.6 1I . . ··f···· .... ····· ..···· . ····················· . ···!
Downloaded by [92.96.139.115] at 01:11 31 January 2016
tions the activity or functional concentration of an ion is nor-
mally derived by taking ionic strength, rather than simply
electrolyte concentration, into account. Accordingly, we ven-
tured a guess that ionic strength (I == 0.51 Cj zj2 where Cj =
molar concentration of the ith ion with valence z) might be a
more appropriate index of salinity stress than either EC or
total electrolyte concentration. This seems to be confirmed by
Figure 5, where the use of ionic strength as a predictor of
growth response eliminates the difference between Na2S04
and NaCI treatments that was evident in Figures 1 and 2. Fur-
ther investigation of ionic strength as an index of salinity
stress is therefore warranted. It must be emphasised that the
common two-phase response to salinity over time (Munns,
2002) makes extrapolation from the current experiment
impossible. Higher proportions of Ca to Na should also be
tested in relation to ionic strength effects.
Because of the convenience of EC as a salinity index we
do not propose that it be replaced routinely but it does seem
as though we can develop a better insight into salinity effects

.Q
that are specific to saIt type (increasingly important in the dis-
~ 1.5 i posal of waste water, for example) by going beyond EC to
g 1.4;
~
ionic strength, which is both an inherent property of salt solu-
1.3 J 1 ..••. Sodium sulphate
tions, affecting the functional concentration (activity) of spe-
--<J- Sodium chloride
1 . 2 L - - - r------·----,c----------,- cific ions, and a more quantitative measure of osmotic
5 10 15 20 potential. This result may have special relevance to wastewa-
Electrical conductivity (dS m·l ) ter irrigation such as that being developed near South African
coalfields, where sulphate-dominated, neutralised mine water
Figure 4 Foliar cation uptake versus electrical conductivity for
is being applied to crops (Annandale et ai., 2002; Jovanovic
Na2S04 and NaCI treatments. Data presented are from experiments
conducted with 900 mg Mn added kg-! dry soil. Error bars depict val- et ai., 2002). Similar wastewaters are common throughout the
ues from the two replicates in each treatment. world wherever acid mine drainage is alleviated through lime
treatment.

4.: ~ Conclusions
Ionic strength appears to be more consistent than electrical
... 3.53 i
...... Sodium sulphate
- 0 - Sodium chloride
conductivity for quantifying the salinity stress imposed by
0 J
irrigated Na2S04 and NaCI salt solutions on kikuyu grass
~
Q.
til
E 2.5 I growing in soil-filled pots. The main difference between the
~
S 2 1 effects of Na2S04 and NaCI on kikuyu nutrition was an
J;! I ......... tt:
Q)
's, 1.~ ~ apparent S04-induced reduction in Ca uptake under Na2S04
~ treatments. This difference was attributed to lower free Ca2+
0
0.5 j activity due to more ion pair formation and a higher ionic
o, strength. Neutral ion pairs, more prevalent in sulphate solu-
0.075 0.125 0.175 0.225 0.275 0.325 0.375
tions, make a contribution to osmotic potential that is not
Ionic Strength
reflected in electrical conductivity measurements. Without
Figure 5 Dry matter yield versus ionic strength for Na2S04 and challenging the routine usefulness of EC, there may neverthe-
NaCI treatments. Data presented are from experiments conducted less be situations in wastewater management where salinity
with 900 mg Mn added kg-! dry soil. Error bars depict values from
thresholds for irrigated crops ought to accommodate ion spe-
the two replicates in each treatment.
cific effects, in which case ionic strength (readily calculated
from routine water analysis) may prove more useful than
Ionic strength asia more consistent index of salinity
electrical conductivity.
stress
Earlier in this paper a discrepancy in growth response to a
Acknowledgements
particular salinity level of sulphate or chloride, as indicated
by EC (Figure 1), was attributed to the failure of EC to cor- The authors extend their thanks to Antoinette Upton (Univer-
rectly reflect both osmotic and ionic interaction effects on sity of Cape Town) for assistance in the laboratory. This work
plant growth. Ionic imbalances may occur through a "swamp- was funded by Shell South Africa (Pty) Ltd, the South Afri-
ing" effect of the dominant ion at root exchange sites produc- can National Research Foundation (Grant number 2047381),
ing either antagonism or synergy and impairing a balanced the Eskom Tertiary Education Support Programme and the
nutrient uptake (Cheeseman, 1988). In dilute electrolyte solu- National Botanical Institute, South Africa.
 S. Afr. J. Plant Soil 2004, 21(2)
References
ANNANDALE, 1. G., JOVANOVIC, N. Z., TANNER, P. D,
BENADE, N, & DU PLESSIS, H. M., 2002. The sustainablility
of irrigation with gypsiferous mine water and implications for the
mining industry in South Africa. Mine Water and the Environ-
ment 21,81-90.
ASHRAF, M., ARF AN, M., & AHMAD, A., 2003. Salt tolerance in
Okra: ion relations and gas exchange characteristics. J. Plant Nut.
26,63-79.
AWADA, S, CAMPBELL, W. F., DUDLEY, L. M., JURINAK, J.
1., & KHAN, M. A., 1995. Interactive effects of sodium chloride,
sodium sulphate, calcium sulphate, and calcium chloride on snap-
bean growth, photosynthesis and ion uptake. J. Plant Nut. 18,
889-900.
AYERS, R. S. & WESTCOT, D. W., 1985. Water Quality for Agri-
culture., Food and Agriculural Organisation of the United
Nations, Rome, Italy.
CAMPBELL, R. B., BOWER, C. A., & RICHARDS, L. A., 1948.
Change of electrical conductivity with temperature and the rela-
tion of osmotic pressure to electrical conductivity and ion con-
centration for soil extracts. Soil Sci. Soc. Am. Proc. 13, 66-69.
CHEESEMAN, J. M., 1988. Mechanisms of salinity tolerance in
Downloaded by [92.96.139.115] at 01:11 31 January 2016
plants. Plant Physiol. 87,547-550.
CURTIN, D., STEPPUHN, H., & SELLES, F., 1993. Plant
responses to sulphate and chloride salinity: growth and ionic rela-
tions. Soil Sci. Soc. Am. J. 57, 1304-1310.
DATTA, K. S., KUMAR, A., VARMA, S. K., & ANGRISH, R.,
1995. Differentiation of chloride and sulphate salinity on the
basis of ionic distribution in genetically diverse cultivars of
wheat. J. Plant Nut. 18,2199-2212.
DEVITT, D. A., 1989. Bermuda grass response to leaching frac-
tions, irrigation salinity and soil types. Agron. J. 8 I, 893-901.
GRATTON, S. R. & GRIEVE, C. M., 1999. Salinity-mineral nutri-
ent relations in horticultural crops. Sci. Hort. 78, 127-157.
HANSEN, E. H. & MUNNS, D. N., 1988. Effect of CaS04 and
NaCI on mineral content of Leucaena leucocephela. Plant Soil
107, 101-105.
JOVANOVIC, N. Z., ANNANDALE, J. G., CLAASSENS, A. S.,
LORENTZ, S. L., TANNER, P. D., AKEN, M. E., & HODG-
SON, F. D. I., 2002. Commercial production of crops irrigated
with gypsiferous mine water. Water SA 28, 413-423.
JANZEN, H. H. & CHANG, C., 1987. Cation nutrition of barley as
influenced by soil solution composition in a saline soil. Can. J.
Soil Sci. 67,616-629.
KA YA, C., AK B.E., & HIGGS, D., 2003. Response of salt-stressed
strawberry plants to supplementary calcium nitrate and/or potas-
sium nitrate. J. Plant Nut. 26, 543-560.
KRAMER, P. J., 1983. Water relations ofplants., Academic Press,
213
New York.
MAAS, E. V. & GRIEVE, c. M., 1987. Sodium induced calcium
deficiency in salt stressed corn. Plant Cell Enviro. 10,559-564.
MANCHANDA, H. R. & SHARMA, S. K., 1989. Tolerance of
chloride and sulphate salinity in chickpea (Cicer arietinum). J.
Agric. Sci. Camb. 113,407-410.
MENGEL, K. & KIRKBY, E. A., 1978. Principles of Plant Nutri-
tion., International Potash Institute, Worblaufen-Bem, Switzer-
land.
MUNNS, R. 2002. Comparative physiology of salt and water stress.
Plant Cell Environ. 25, 239-250.
PAEK, K. Y., CHANDLER, S. F., & THORPE, T. A., 1988. Physio-
logical effects ofNa2S04 and NaCI on callus cultures of Brassica
campestri (Chinese Cabbage). Physio/. Plant. 72, 160-166.
PAPADOPOULOS, I., 1986. Effect of high sulphate irrigation
waters on soil salinity and yields. Agron. J. 78, 429-432.
PARKHURST, D. L. & APPELO, C. A. J., 1999. User's guide to
PREEQC (version 2) - a computer program for speciation, batch-
reaction, one-dimensional transport, and inverse geochemical cal-
culations. U.S. Geological Survey, Denver, Colorado.
PEACOCK, C. H., DUDECK, A. E., & WILDMON, J. C., 1993.
Growth and mineral content of St. Augustine grass cultivars in
response to salinity. J. Am. Soc. Hort. Sci. 118,464-469.
RA YMENT, G. E. & VERRALL, K. A., 1980. Soil manganese tests
and the comparative tolerance of kikuyu and white clover to man-
ganesetoxicity. Trop. Grass. 14,105-114.
RHOADES, J. D., 1982. Soluble Salts. In Page, A. L, Miller, R. H,
and Keeney, D. R. (ed.). Methods of Soil AnalYSis. Part 2. Chem-
ical and Microbiological Properties. Soil Science Society of
America, Inc., Madison, Wisconsin.
RUSSELL, J. S., 1976. Comparative salt tolerance of some tropical
and temperate legumes and tropical grasses. Aus. J. Exp. Agric.
Anim. Husb. 16, 103-109.
SNOEYINK, V. L. & JENKINS, D., 1980. Water Chemistry, John
Wiley & Sons, New York.
SPOSITO, G., 1989. The Chemistry of Soils., Oxford University
Press, Oxford.
SUHAYDA, C. G., REDMANN, R. E., HARVEY, B. L., & CIPY-
WNYK, A. L., 1992. Comparative response of cultivated and
wild barley species to salinity stress and calcium supply. Crop
Sci. 32, 154-163.
WARNE, P., GUY, R. D., ROLLINS, L., & REID, D. M., 1990. The
effects of sodium sulphate and sodium chloride on growth, mor-
phology, photosynthesis and water use efficiency of Chenopo-
dium rubrum. Can. J. Bot. 68, 999-1006.
YEO, A. R., 1983. Salinity resistance: physiologies and price. Phys-
io/. Plant 58,214-222.