Plant Soil (2010) 326:381–391

DOI 10.1007/s11104-009-0019-9

REGULAR ARTICLE

Alleviation of detrimental effects of NaCl by silicon nutrition

in salt-sensitive and salt-tolerant genotypes of sugarcane
(Saccharum officinarum L.)
M. Ashraf & Rahmatullah & M. Afzal & R. Ahmed &
F. Mujeeb & A. Sarwar & L. Ali

Received: 30 September 2008 / Accepted: 28 April 2009 / Published online: 19 May 2009
# Springer Science + Business Media B.V. 2009

Abstract Salt stress is a major environmental factor
which adversely affects the crop yield and quality.
However, adequate regulation of mineral nutrients
may ameliorate the deleterious effects of salts and
help to sustain crop productivity under salt stress.
Salt-sensitive (SPF 213) and salt-tolerant (HSF 240)
sugarcane genotypes were grown in gravel at 0 and
100 mM NaCl by supplying 0, 1.4 mM, 2.1 mM and
2.8 mM of Si as calcium silicate. Results revealed that
plants treated with NaCl alone showed a significant
(P≤0.05) reduction in dry matter production, K+
concentration, cane yield and juice quality in both
Responsible Editor: Yong Chao Liang.
M. Ashraf (*) : M. Afzal
University College of Agriculture,
University of Sargodha,
Sargodha, Pakistan
e-mail: mashraf_pk94@hotmail.com
genotypes but the magnitude of reduction was
relatively more in salt-sensitive genotype than salt-
tolerant. Addition of Si significantly (P≤0.05) re-
duced the uptake and translocation of Na+ but
increased K+ concentrations particularly in shoots of
both sugarcane genotypes. Cane yield and yield
attributes were significantly (P≤0.05) higher where
Si was added. Juice quality characteristics were
significantly (P≤0.05) improved in salt-sensitive and
salt-tolerant sugarcane genotypes with the application
of Si. The results suggested that added Si interacted
with Na+, reduced its uptake and transport to shoots
and consequently improved cane yield and juice
quality in salt-sensitive and salt-tolerant sugarcane
genotypes under salt stress.
Keywords Genotype . Juice quality . Salt tolerance .
Silicon . Sugarcane

Rahmatullah : F. Mujeeb : L. Ali

Institute of Soil & Environmental Sciences, Introduction
University of Agriculture,
Faisalabad, Pakistan Long considered by plant physiologist as a non-
essential nutrient, the importance of silicon (Si)
R. Ahmed
Department of Crop Physiology, University of Agriculture, has only recently been recognized (Epstein 1999;
Faisalabad, Pakistan Richmond and Sussman 2003; Liang et al. 2007).
Silicon is the second most abundant mineral element
A. Sarwar
in the soil after oxygen and comprises 28% of the
Sugarcane Research Institute,
Ayub Agricultural Research Institute, earth crust (Gong et al. 2006). Silicon is taken up by
Faisalabad, Pakistan plants in the form of silicic acid [Si(OH)4], an
382
uncharged monomeric molecule, when the solution
pH is below 9 (Ma et al. 2006). In shoot, silicic acid is
concentrated through the loss of water (transpiration)
and is polymerized to form silica gel (SiO2.nH2O)
(Raven 2003). The process of Si polymerization
converts silicic acid to colloidal silicic acid and
finally to silica gel with increasing silicic acid
concentration (Ma and Takahashi 1993).
Silicon is deposited as a 2.5 µm layer in the space
immediately beneath the thin cuticle layer, forming a
cuticle-Si double layer in leaf blades (Neumann and
Nieden 2001). Most of the beneficial effects of Si are
attributed to its deposition in cell walls of the roots,
leaves and stems (Ma et al. 2006). For example,
deposition of Si in roots reduces apoplastic bypass
flow and provides binding sites for salts resulting in
reduced uptake and translocation of salts from roots to
shoots. According to Bradbury and Ahmad (1990)
and Ahmad et al. (1992) silica deposition in the leaves
limits transpiration and hence salt accumulation in
wheat. Romero-Aranda et al. (2006) have suggested
that silicate crystals deposited in the epidermal cells
form a barrier that reduces water loss through cuticle,
which in turn, contributes to salt dilution, mitigating
salt toxicity effects in tomato. More recently, Si
benefits on salt tolerance of barley and cucumber
have been related to antioxidant enzyme activity
(Liang et al. 2003; Zhu et al. 2004).
Past studies have mostly addressed the role of Si to
alleviate salt-stress in wheat (Ahmad et al. 1992;
Liang et al. 2003, 2005), tomato (Romero-Aranda et
al. 2006), rice (Yeo et al. 1999), cucumber (Zhu et al.
2004), barley (Liang 1999), maize (Shu and Liu
2001) and alfalfa (Wang and Han 2007) but very little
is known about the beneficial effects of Si on salt-
tolerance of sugarcane which is moderately sensitive
to salinity (Shannon 1997) with a threshold for yield
reduction at 1.7 dS m−1 (Maas and Grieve 1990). The
present study is therefore, designed to investigate Si-
mediated alleviation of detrimental effects of NaCl in
sugarcane genotypes differing in salt-tolerance.
Material and methods
Plant material
Two sugarcane genotypes, one of them (SPF 213)
identified as salt-sensitive and other (HSF 240) as
Plant Soil (2010) 326:381–391
salt-tolerant (Ashraf et al. 2007) were grown in gravel
to study the role of Si in alleviating salt stress in
sugarcane. Two single nodded stem sections (5 cm
long) of each genotype were planted in plastic pot
filled with 55 kg of pre-washed gravel. These pots
were irrigated daily with tap water to keep the gravel
moist.
Basal nutrient solution
Ammonium sulfate [(NH4)2SO4], potassium nitrate
(KNO 3 ), ammonium dihydrogen phosphate
(NH4H2PO4), calcium nitrate [Ca(NO3)2] and mag-
nesium sulfate (MgSO4.7H2O) were used to add
8 mM N, 1 mM P, 3 mM K, 2 mM Ca, 0.5 mM Mg
and 0.5 mM S to the nutrient solution. Different
micro-elements included 25 µM B (H3BO3), 2 µM
Mn (MnSO4), 2 µM Zn (ZnSO4), 0.5 µM Mo
(H2MoO4) and 50 µM Fe (Fe-EDTA) were also
added to nutrient solution (Johnson et al. 1957).
Demineralized water was used to prepare solutions of
various nutrients.
Silicon and NaCl treatments
After 20 days of planting, two NaCl levels: 0 (no
added NaCl) and 100 mM NaCl were imposed in
triplicate to plants supplied with four levels of Si: viz.
0 (no added Si), 1.4 mM Si, 2.1 mM Si and 2.8 mM
Si, making eight combination treatments. Calcium
silicate after digesting with 50% H2O2 and 50%
NaOH was used to maintain the Si levels. Calcium in
the nutrient solution was balanced among all treat-
ments including control by subtracting additional Ca
introduced by calcium silicate from calcium nitrate
and then adding balanced Ca by calcium nitrate. The
initial pH of nutrient solution after the addition of
different levels of Si was 8.0–9.5 which was adjusted
to 6.0–6.5 using H2SO4 before applying to plants. Salt
level was maintained in the respective pots by
providing saline water of EC 10 dS m−1 at regular
intervals while tap water was used in control plants.
Ionic relations
After 80 days of planting, one plant from each pot
was harvested while second was allowed to mature.
The harvested plants were separated into roots and
shoots, dried to a constant weight at 70°C in a forced-
Plant Soil (2010) 326:381–391
air oven (EYELA, WFO-600ND) and weighed to
obtain dry matter yield. The samples were then
ground with a Wiley mill fitted with a stainless steel
chamber and blades (IKA-WERKE, GMBH & CO.
KG, Germany). The fine ground samples of roots and
shoots were digested with di-acid mixture of HNO3
and HClO4 (3:1) for K+ and Na+ analysis (Yoshida et
al. 1976). The concentrations of K+ and Na+ in the
digest were estimated by flamephotometery (Jenway
PFP 7, ELE Instrument Co. Ltd.). For Si estimation,
0.1 g ground samples of roots and shoots were
digested in 2 ml of 50% H2O2 and 4 ml of 50%
NaOH for 2 h at 150°C. Silicon was estimated by
molybdatevanadate blue color method using UV
spectrophotometer (Shimdzu, Spectron 100, Japan)
at 650 nm (Elliot and Snyder 1991).
Yield and yield-attributes
At maturity (350 day old crop), second plant from each
pot was harvested. Cane yield and yield contributing
parameters like number of canes per plant, cane height
and internode’s distance were recorded after topping and
stripping the cane of trash.
Juice quality
Cane samples were crushed through a three-roller
miller. The juice samples were analyzed for Brix (%
soluble solids in juice) by a brix refractrometer. Pol
(% sucrose in juice) was determined by an automatic
sucrolyser after clarifying the juice with lead acetate.
Percent commercial cane sugar (%CCS) was calcu-
lated on the basis of following formula (Anonymous
1970):
   
3P Fþ5 B Fþ3
%CCS ¼ 1

1

2 100 2 100
P Pol%
F Fiber% cane (CCS was calculated at constant
fiber value of 12.5%)
B Brix%
Sugar recovery was calculated as follows:
Sugar recovery ¼ %CCS  0:94
383
Statistical analysis
The data were subjected to statistical analysis using
computer software MSTAT-C (Russell and Eisenmith
1983) and following methods described by Gomez
and Gomez (1984). Completely randomized design
(CRD) was employed for analysis of variance and
Duncan’s multiple range tests were used to separate
treatment means at P≤0.05.
Results
Plant growth
A significant (P≤0.05) main and interactive effect of
NaCl, Si and genotypes on plant growth occurred in
sugarcane (Table 1). Applied NaCl reduced shoot dry
matter by 46% and root dry matter by 48% in salt-
sensitive genotype (SPF 213) while in salt-tolerant
genotype (HSF 240) the reduction was 25% and 29%
respectively, as compared to control. Addition of
different levels of Si to the growth medium interacted
with Na+, reduced its uptake and translocation to
shoots and consequently improved plant growth in
both genotypes. Although all levels of applied Si were
effective to mitigate the adverse effects of NaCl on
plant growth but maximum increase in plant growth
was observed at highest rate of Si application
(2.8 mM). Moreover, the beneficial effects of added
Si were more marked in salt-sensitive genotype than
salt-tolerant, particularly in the presence of NaCl.
Ionic relations
A significant (P≤0.05) main and interactive effect of
NaCl, Si and genotypes on K+, Na+ and Si concen-
trations occurred in sugarcane. Potassium concentra-
tion in shoots and roots of salt-sensitive (SPF 213)
and salt-tolerant (HSF 240) sugarcane genotypes was
markedly reduced in the presence of NaCl as
compared to control. Addition of different levels of
Si increased shoot K+ concentration by 10 to 54% and
root K concentration by 15 to 37% in salt-sensitive
genotype; while in salt-tolerant genotype the increase
was 3 to 11% in shoot K+ and 2 to 18% in root K+
concentrations as compared to salt-stressed plants
without added Si (Table 2). Sodium concentration in
plant tissues was significantly (P≤0.05) higher in the
384 Plant Soil (2010) 326:381–391

Table 1 Shoot and root dry matter of salt-sensitive and salt-tolerant sugarcane genotypes grown under salt stress by supplying Si
(Figures are the means of 3 replications)

Treatments Shoot dry matter (g plant−1) Root dry matter (g plant−1)

SPF 213 HSF 240 SPF 213 HSF 240

(Salt-sensitive) (Salt-tolerant) (Salt-sensitive) (Salt-tolerant)

Control 77.2 cdea 86.4 b–e 5.0 ef 7.6 def

NaCl 41.4 f 64.8 ef 2.6 g 5.4 ef
Si1 86.0 b–e 96.3 bcd 5.9 ef 9.8 c–f
Si1 + NaCl 64.3 ef 75.1 de 8.4 c–f 8.3 c–f
Si2 89.1 b–e 101.7 abc 6.3 def 11.9 cde
Si2 + NaCl 76.9 cde 80.2 cde 8.5 c–f 12.6 cd
Si3 107.5 ab 124.8 a 7.0 def 19.6 a
Si3 + NaCl 97.4 bcd 100.3 bcd 17.1 ab 14.9 bc
LSD(0.05) 22.25 5.80
(Salinity × Si × Genotype)
a
Mean values followed by the same letter in each column are not significantly different at P≤0.05. Control, nutrient solution without
added Si; NaCl, 100 mM NaCl; Si1, 1.4 mM Si; Si1 + NaCl, 1.4 mM Si + 100 mM NaCl; Si2, 2.1 mM Si; Si2 + NaCl, 2.1 mM Si +
100 mM NaCl; Si3, 2.8 mM Si; Si3 + NaCl, 2.8 mM Si + 100 mM NaCl

presence of NaCl and added Si reduced Na+ uptake
and transport to shoots (Table 3) with a resultant
increase in K+/Na+ ratio in shoot of both genotypes
(data not reported). The ratio of shoot Na+/root Na+
was significantly (P≤0.05) reduced in salt-stressed
plants of both genotypes by various levels of added
Si, indicating Si-mediated reduction in the transport
of Na+ from roots to shoots (Fig. 1). Silicon
concentration in shoots and roots of both genotypes
was significantly (P≤0.05) increased with increasing
Si levels in the growth medium, regardless of NaCl
treatment. However, the increase was more prominent
in salt-sensitive genotype than salt-tolerant genotype
(Table 4). Concentrations of Si in sugarcane shoots

Table 2 Potassium concentration of salt-sensitive and salt-tolerant sugarcane genotypes grown under salt stress by supplying Si
(Figures are the means of 3 replications)

Treatments Concentration in shoot (mg g−1) Concentration in root (mg g−1)

SPF 213 HSF 240 SPF 213 HSF 240

(Salt-sensitive) (Salt-tolerant) (Salt-sensitive) (Salt-tolerant)

Control 33.25 cdea 55.50 ab 50.33 ab 46.13 ab

NaCl 26.51 e 45.79 a–e 33.77 b 36.96 ab
Si1 40.12 b–e 58.04 ab 57.74 ab 51.21 ab
Si1 + NaCl 29.13 de 47.15 a–e 38.88 ab 37.82 ab
Si2 42.22 b–e 59.26 ab 59.16 a 57.74 ab
Si2 + NaCl 32.81 cde 48.80 a–d 44.21 ab 37.97 ab
Si3 53.46 abc 68.53 a 62.72 ab 66.14 a
Si3 + NaCl 40.83 b–e 50.83 a–d 46.22 ab 43.62 ab
LSD(0.05) 18.58 26.49
(Salinity × Si × Genotype)
a
Mean values followed by the same letter in each column are not significantly different at P≤0.05. Control, nutrient solution without
added Si; NaCl, 100 mM NaCl; Si1, 1.4 mM Si; Si1 + NaCl, 1.4 mM Si + 100 mM NaCl; Si2, 2.1 mM Si; Si2 + NaCl, 2.1 mM Si +
100 mM NaCl; Si3, 2.8 mM Si; Si3 + NaCl, 2.8 mM Si + 100 mM NaCl
Plant Soil (2010) 326:381–391 385

Table 3 Sodium concentration of salt-sensitive and salt-tolerant sugarcane genotypes grown under salt stress by supplying Si (Figures
are the means of 3 replications)

Treatments Concentration in shoot (mg g−1) Concentration in root (mg g−1)

SPF 213 HSF 240 SPF 213 HSF 240

(Salt-sensitive) (Salt-tolerant) (Salt-sensitive) (Salt-tolerant)

Control 14.47 hia 11.63 i 9.54 j 10.67 ij

NaCl 61.80 a 42.10 c 28.51 a 21.79 cd
Si1 18.67 ghi 13.93 hi 11.03 hij 11.60 g–j
Si1 + NaCl 52.30 b 35.13 cde 23.28 c 25.98 b
Si2 20.17 ghi 15.30 hi 12.48 fgh 12.7 f–i
Si2 + NaCl 38.50 cd 34.34 cde 21.06 d 28.98 a
Si3 22.17 fgh 16.27 hi 13.65 f 13.11 fg
Si3 + NaCl 30.47 def 27.17 efg 18.97 e 30.07 a
LSD(0.05) 8.06 1.60
(Salinity × Si × Genotype)
a
Mean values followed by the same letter in each column are not significantly different at P≤0.05. Control, nutrient solution without
added Si; NaCl, 100 mM NaCl; Si1, 1.4 mM Si; Si1 + NaCl, 1.4 mM Si + 100 mM NaCl; Si2, 2.1 mM Si; Si2 + NaCl, 2.1 mM Si +
100 mM NaCl; Si3, 2.8 mM Si; Si3 + NaCl, 2.8 mM Si + 100 mM NaCl

determined for various treatments in the presence of NaCl, but application of different levels of Si signif-
NaCl were positively correlated with shoot dry matter icantly (P≤0.05) improved these growth traits in salt-
of salt-sensitive (R2 =0.99) and salt-tolerant (R2 =0.89) sensitive and salt-tolerant sugarcane genotypes as
genotypes but negatively correlated with Na+ concen- compared to salt-stressed plants without Si (Table 6).
tration in shoots of salt-sensitive (R2 =0.95) and of
salt-tolerant (R2 =0.97) genotypes (Figs. 2 & 3).
2.5
Yield and yield attributes a
ab
A significant (P≤0.05) main and interactive effect of 2 abc bcd
Shoot Na+/root Na+ ratio

NaCl, Si and genotypes on cane yield and yield cd

attributes occurred in sugarcane. Cane yield was
reduced by 35% in salt-sensitive genotype (SPF 1.5 de
213) and 28% in salt-tolerant genotype in the presence def
of NaCl as compared to control. Addition of Si to the 1 fg
saline growth medium significantly (P≤0.05) mitigated
the deleterious effects of NaCl and improved cane
yield in salt-sensitive genotype by 20% with 1.4 mM 0.5 Salt-sensitive genotype (SPF 213)
Si, 29% with 2.1 mM Si and 77% with 2.8 mM Si;
Salt-tolerant genotype (HSF 240)
while in salt-tolerant genotype the increase was 10%,
0
15% and 22% with 1.4, 2.1 and 2.8 mM Si
NaCl NaCl+Si1 NaCl+Si2 NaCl+Si3
respectively, as compared to salt-stressed plants with-
Treatments
out Si. Addition of different levels of Si significantly
(P≤0.05) improved internode’s distance in both sugar- Fig. 1 Shoot Na+/root Na+ ratio of salt-sensitive and salt-
cane genotypes; however, the magnitude of increment tolerant sugarcane genotypes grown under salt stress by
was relatively higher in salt-sensitive genotype than supplying Si (Values are the means of 3 replications; LSD0.05 =
0.42). NaCl, 100 mM NaCl; NaCl+Si1, 100 mM NaCl+1.4 mM
salt-tolerant (Table 5). Numbers of canes plant−1 and Si; NaCl+ Si2, 100 mM NaCl+2.1 mM Si; NaCl+Si3, 100 mM
cane height were slightly reduced in the presence of NaCl+2.8 mM Si
386 Plant Soil (2010) 326:381–391

Table 4 Silicon concentration of salt-sensitive and salt-tolerant sugarcane genotypes grown under salt stress by supplying Si (Figures
are the means of 3 replications)

Treatments Concentration in shoot (mg g−1) Concentration in root (mg g−1)

SPF 213 HSF 240 SPF 213 HSF 240

(Salt-sensitive) (Salt-tolerant) (Salt-sensitive) (Salt-tolerant)

Control 9.1 la 4.5 m 5.4 e 7.2 e

NaCl 13.5 k 6.2 m 5.7 e 4.9 e
Si1 33.2 i 30.2 j 13.9 d 17.6 d
Si1 + NaCl 39.8 gh 37.5 h 14.7 d 16.0 d
Si2 49.2 e 42.4 fg 27.3 c 26.7 c
Si2 + NaCl 52.2 d 44.7 f 24.9 c 32.4 b
Si3 61.5 c 59.8 c 32.9 b 43.4 a
Si3 + NaCl 69.4 a 64.9 b 32.3 b 45.0 a
LSD(0.05) 2.77 4.38
(Salinity × Si × Genotype)
a
Mean values followed by the same letter in each column are not significantly different at P≤0.05. Control, nutrient solution without
added Si; NaCl, 100 mM NaCl; Si1, 1.4 mM Si; Si1 + NaCl, 1.4 mM Si + 100 mM NaCl; Si2, 2.1 mM Si; Si2 + NaCl, 2.1 mM Si +
100 mM NaCl; Si3, 2.8 mM Si; Si3 + NaCl, 2.8 mM Si + 100 mM NaCl

Juice quality sucrose in juice) due to applied NaCl was slightly

higher in salt-sensitive genotype than salt-tolerant.
A significant (P≤0.05) main and interactive effect of Addition of Si to the growth medium significantly
NaCl, Si and genotypes on juice quality occurred in (P≤0.05) improved Pol in both genotypes as com-
sugarcane. Applied NaCl reduced Brix (% soluble pared to Si-untreated plants (Fig. 5). Applied NaCl
solids in juice) in salt-sensitive genotype by 28% and reduced commercial cane sugar (CCS) of salt-
in salt-tolerant genotype by 24% as compared to sensitive genotype by 30% and salt-tolerant genotype
control. Addition of different levels of Si significantly by 26% as compared to control. Addition of Si
(P≤0.05) improved this juice quality parameter in improved CCS in both salt-sensitive and salt-tolerant
both genotypes (Fig. 4). The reduction in Pol (% sugarcane genotypes as compared to Si-untreated

70 120
y = -0.5822x + 71.205
R2 = 0.95
Na+ concentration in shoot (mg g-1)

100
56 y = 0.580x+57.85
Shoot dry matter (g plnat-1)

R2 = 0.89
80
42
60
y = -0.2477x + 44.167
28 R2 = 0.97 y = 0.993x+26.57
40 R2 = 0.99

14 Salt-sensitive genotype (SPF 213) Salt-sensitive genotype (SPF 213)

20
Salt-tolerant genotype (HSF 240) Salt-tolerant genotype (HSF 240)
0 0
0 20 40 60 80 0 20 40 60 80
Si concentration in shoot (mg g-1) Silicon concentration in shoot (mg g-1)

Fig. 2 Relationship between Si concentration in sugarcane Fig. 3 Relationship between Si concentration in sugarcane
shoot with shoot Na+ concentration in salt-sensitive and salt- shoot with shoot dry matter in salt-sensitive and salt-tolerant
tolerant sugarcane genotypes grown under salt stress by sugarcane genotypes grown under salt stress by supplying
supplying different levels of Si different levels of Si
Plant Soil (2010) 326:381–391 387

Table 5 Cane yield and internode’s distance of salt-sensitive and salt-tolerant sugarcane genotypes grown under salt stress by
supplying Si (Figures are the means of 3 replications)

Treatments Cane yield (kg plant−1) Internode’s distance(cm)

SPF 213 HSF 240 SPF 213 HSF 240

(Salt-sensitive) (Salt-tolerant) (Salt-sensitive) (Salt-tolerant)

Control 6.11 cdea 6.69 a–d 10.80 def 11.27 cde

NaCl 3.95 g 5.23 f 9.15 f 9.75 ef
Si1 6.50 a–d 7.38 ab 11.87 bcd 11.95 bcd
Si1 + NaCl 4.74 f 5.78 ef 12.11 bcd 11.17 cde
Si2 6.98 abc 7.68 ab 12.39 bcd 12.55 bcd
Si2 + NaCl 5.09 f 6.03 de 12.44 bcd 12.43 bcd
Si3 7.21 ab 8.83 a 13.22 abc 14.63 a
Si3 + NaCl 6.98 ab 6.38 bc 13.49 ab 12.95 abc
LSD(0.05) 0.65 1.81
(Salinity × Si × Genotype)
a
Mean values followed by the same letter in each column are not significantly different at P≤0.05. Control, nutrient solution without
added Si; NaCl, 100 mM NaCl; Si1, 1.4 mM Si; Si1 + NaCl, 1.4 mM Si + 100 mM NaCl; Si2, 2.1 mM Si; Si2 + NaCl, 2.1 mM Si +
100 mM NaCl; Si3, 2.8 mM Si; Si3 + NaCl, 2.8 mM Si + 100 mM NaCl

plants (Fig. 6). Sugar recovery was also significantly Discussion

(P≤0.05) reduced in salt-stressed plants of both
sugarcane genotypes but addition of different levels
of Si mitigated the deleterious effects of NaCl and
improved sugar recovery by 16 to 43% in salt-
sensitive genotype and 18 to 53% in salt-tolerant
genotype as compared to Si-untreated plants (Fig. 7).
Salt stress is posing a severe threat to increasing
demand of food in many arid and semiarid regions of
the world (Benlloch et al. 1994; Akram et al. 2002;
Wahid 2004). Different techniques including engi-
neering and reclamation approaches, use of salt-

Table 6 Number of canes plant−1 and cane height of salt-sensitive and salt-tolerant sugarcane genotypes grown under salt stress by
supplying Si (Figures are the means of 3 replications)

Treatments Number of canes plant−1 Cane height (m)

SPF 213 HSF 240 SPF 213 HSF 240

(Salt-sensitive) (Salt-tolerant) (Salt-sensitive) (Salt-tolerant)

Control 6.0 cdea 6.66 b–e 1.70 bcd 1.72 a–d

NaCl 4.0 e 5.33 de 1.46 d 1.66 cd
Si1 6.67 b–e 7.33 a–d 1.93 abc 1.84 a–d
Si1 + NaCl 5.33 de 6.66 b–e 1.76 a–d 1.71 b–d
Si2 7.33 a–d 9.0 ab 1.99 abc 1.87 abc
Si2 + NaCl 6.67 b–e 7.0 a–e 1.90 abc 1.80 a–d
Si3 8.0 abc 9.33 a 2.13 a 2.01 abc
Si3 + NaCl 7.0 a–e 7.33 a–d 2.12 ab 1.95 abc
LSD(0.05) 1.38 0.35
(Salinity × Si × Genotype)
a
Mean values followed by the same letter in each column are not significantly different at P≤0.05. Control, nutrient solution without
added Si; NaCl, 100 mM NaCl; Si1, 1.4 mM Si; Si1 + NaCl, 1.4 mM Si + 100 mM NaCl; Si2, 2.1 mM Si; Si2 + NaCl, 2.1 mM Si +
100 mM NaCl; Si3, 2.8 mM Si; Si3 + NaCl, 2.8 mM Si + 100 mM NaCl
388 Plant Soil (2010) 326:381–391

Salt-sensitive genotype (SPF 213) Salt-sensitive genotype (SPF 213)

Commercial cane sugar (%)

20
Salt-tolerant genotype (HSF 240) Salt-tolerant genotype (HSF 240)
Brix (% soluble solids in juice)

25 ab a 16 a
bc b b
cd cde c c c
ef def efg d
20 fg gh
12 d d
h e e e
f
i g
jk jk ij 8 h
15 k
4
10
0
5

l
1

3
aC
ro

aC

aC

aC
Si

Si

Si
nt

N
Co

1+

2+

3+
Si

Si

Si
0
Treatments
l

3
l

l
ro

aC

aC

aC
aC
Si

Si

Si
nt

N
Co

1+

2+

3+
Fig. 6 Commercial cane sugar (the percentage by weight of the
Si

Si

Si
Treatments quantity of cane which would be recovered as pure sucrose) of
salt-sensitive and salt-tolerant sugarcane genotypes grown
Fig. 4 Brix (% soluble solids in juice) of salt-sensitive and under salt stress by supplying Si (Values are the means of 3
salt-tolerant sugarcane genotypes grown under salt stress by replications; LSD0.05 = 0.62). Control, nutrient solution without
supplying Si (Values are the means of 3 replications; LSD0.05 = added Si; NaCl, 100 mM NaCl; Si1, 1.4 mM Si; Si1+NaCl,
1.28). Control, nutrient solution without added Si; NaCl, 1.4 mM Si+100 mM NaCl; Si2, 2.1 mM Si; Si2+NaCl, 2.1 mM
100 mM NaCl; Si 1 , 1.4 mM Si; Si 1 +NaCl, 1.4 mM Si+100 mM NaCl; Si3, 2.8 mM Si; Si3+NaCl, 2.8 mM
Si+100 mM NaCl; Si2, 2.1 mM Si; Si2+NaCl, 2.1 mM Si+100 mM NaCl
Si+100 mM NaCl; Si3, 2.8 mM Si; Si3+NaCl, 2.8 mM Si+
100 mM NaCl
indicator to assess plant tolerance to salinity. Similar
responses of different crop species to NaCl were
tolerant genotypes and adequate regulation of mineral earlier reported (Cachorro et al. 1994; Perez-Alfocea
nutrients may help to sustain the productivity of salt et al. 1996; Liang 1999; Al-Aghabary et al. 2004;
affected soils (Asch et al. 1999; Akhtar et al. 2001; Gunes et al. 2007). These authors suggested that one
Mahar et al. 2003; Ashraf et al. 2007). In present of the primary plant responses to NaCl was the
study, NaCl-induced reduction in cane yield and juice decrease in K+ concentration in plant tissues and thus
quality in salt-sensitive and salt-tolerant sugarcane the substitution of K+ by Na+ in saline soil resulted in
genotypes was ascribed to increased absorption of low K+/Na+ ratio which damaged the plant metabo-
Na+ which had inverse relationship with K+ and lism and ultimately yield and yield attributes were
leading to reduced K+/Na+ ratio which is a good severely affected. Both sugarcane genotypes behaved

Salt-sensitive genotype (SPF 213)

Salt-sensitive genotype (SPF 213) Salt-tolerant genotype (HSF 240)
25 15 a
Salt-tolerant genotype (HSF 240)
Pol (% sucrose in juice)

Sugar recovery (%)

b b
a c c c
12 d
20 b
c d d
cd de def e e e
efg fg 9 fg f
hi gh g
15 ij jk h
kl kl
m
l 6
10
3
5
0
l

0
ro

aC

aC

aC

aC
Si

Si

Si
nt

N
Co

1+

2+

3+
l

l
l

l
ro

aC

aC
aC

aC
Si

Si

Si

Si

Si

Si
nt

N
Co

1+

2+

3+

Treatments
Si

Si

Si

Treatments
Fig. 7 Sugar recovery (the sugar bagged per unit of cane
Fig. 5 Pol (% sucrose in juice) of salt-sensitive and salt-tolerant crushed) of salt-sensitive and salt-tolerant sugarcane genotypes
sugarcane genotypes grown under salt stress by supplying Si grown under salt stress by supplying Si (Values are the means
(Values are the means of 3 replications; LSD0.05 = 0.95). of 3 replications; LSD0.05 = 0.60). Control, nutrient solution
Control, nutrient solution without added Si; NaCl, 100 mM without added Si; NaCl, 100 mM NaCl; Si1, 1.4 mM Si;
NaCl; Si1, 1.4 mM Si; Si1+NaCl, 1.4 mM Si+100 mM NaCl; Si1+NaCl, 1.4 mM Si+100 mM NaCl; Si2, 2.1 mM Si;
Si2, 2.1 mM Si; Si2+NaCl, 2.1 mM Si+100 mM NaCl; Si3, Si 2 +NaCl, 2.1 mM Si+100 mM NaCl; Si 3 , 2.8 mM
2.8 mM Si; Si3+NaCl, 2.8 mM Si+100 mM NaCl Si; Si3+NaCl, 2.8 mM Si+100 mM NaCl
Plant Soil (2010) 326:381–391
differentially to NaCl depending on their capability to
selectively absorb K+ over Na+. Better performance of
salt-tolerant genotype (HSF 240) in the presence of
NaCl was due to its lower accumulation of Na+ than
salt-sensitive genotype (SPF 213). Munns and James
(2003) also demonstrated that genotype with lowest
Na+ concentration produced greatest biomass and vice
versa. Lingle and Wiegand (1997) reported that
applied NaCl had a profound effect on the biosynthe-
sis of sucrose in the leaf and its translocation to stalk
for storage. According to Tazuke and Wada (2002),
NaCl modulated the activities of sugar metabolizing
enzymes in a number of crops. It reduced the
activities of sucrose synthase and starch phosphory-
lase and enhanced those of acid and neutral inver-
tases. Addition of different levels of Si under salt
stress interacted with Na+, reduced its uptake and
transport to shoots with a resultant improvement in
cane yield and juice quality of both genotypes. The
ameliorative effect of added Si in alleviating delete-
rious effects of NaCl could be related to Si being
irreversibly precipitated as amorphous silica (SiO2.
nH2O) in cell walls and lumens and thus reduced the
translocation of salts to shoots (Bradbury and Ahmad
1990; Epstein 1999; Gong et al. 2006; Gunes et al.
2007). Yeo et al. (1999) also demonstrated that the
deposition and polymerization of silicate in the
endodermis and rhizodermis might offer possible
mechanism by which Si could physically block the
transpirational bypass flow across the roots and
restrained the apoplastic transport of salts.
Furthermore, a marked reduction in the ratio of
shoot Na+ to root Na+ of both sugarcane genotypes
with the supplementation of Si in the presence of
NaCl also suggested that Na+ uptake and transporta-
tion into shoots from the roots was greatly inhibited
by added Si under salt stress. In experiments with rice
(Matoh et al. 1986), wheat (Ahmad et al. 1992),
barley (Liang 1999) and alfalfa (Wang and Han
2007), it was demonstrated that added Si to the
nutrient solution diminished Na+ contents in the
shoots and thereby reducing the ratio of shoot Na+
to root Na+. It was interesting to note that HSF 240
(salt-tolerant genotype) retained more Na+ than SPF
213 (salt-sensitive genotype) in roots in the presence
of Si which suggested that transportation of Na+ from
roots to shoots was an important physiological
contribution to salt-tolerance. Liang (1999) also
reported a lower ratio of shoot Na+ to root Na+ for
389
salt-tolerant cultivar compared to salt-sensitive, indi-
cating that Na+ exclusion from the shoots was an
important mechanism in salt-tolerance. Added Si was
also found to increase K+ concentration particularly in
the shoots of both genotypes. Potassium played an
important role in contributing to the survival of crop
plants under salt stress (Cakmak 2005) and the
possible mechanism causing Si to stimulate the plants
to absorb K+ under salt stress was the activation of
H+-ATPase in the membranes (Liang 1999). Accord-
ing to Savant et al. (1999) the improvement in yield
and juice quality of both sugarcane genotypes by Si
under salt stress could be attributed to Si-enzyme
complexes which served as a protector or regulator of
photosynthesis and enzyme activity in the presence of
NaCl.
The interaction between Si and NaCl varied with
plant species and varieties (Liang 1999; Gunes et al.
2007). In present study, the ameliorative effects of
added Si were more pronounced in salt-sensitive
genotype than salt-tolerant. Similar results were
observed in experiments with wheat (Ahmad et al.
1992) and rice genotypes (Yeo et al. 1999) who
reported that salt-tolerant genotypes were least re-
sponsive to Si application under salt stress. These
authors further suggested that differences in the
response of genotypes to Si could be related to the
size of bypass flow and/or the properties that affected
the polymerization of silicate.
Conclusion
The results confirmed that applied NaCl markedly
increased Na+ concentration in plant tissue which
adversely affected the uptake of K+, cane yield and
juice quality in both salt-sensitive and salt-tolerant
sugarcane genotypes. Addition of different levels of
Si to growth medium interacted with Na+, reduced its
uptake and transport to shoots and ultimately im-
proved cane yield and juice quality in both genotypes.
Although all levels of added Si were effective to
mitigate the adverse effects of NaCl but best results
were found when Si was applied @ 2.8 mM. In this
study, Si-induced salt-tolerance in sugarcane genotypes
was believed to be associated with decreased Na+
concentration and increased K+ concentration particu-
larly in shoots with a resultant improvement in shoot
K+/Na+ ratio of both genotypes.
390
Acknowledgements This research is financially supported by
Higher Education Commission of Pakistan through Indigenous
Ph.D. Scholarship Scheme. The authors are grateful to Director,
Sugarcane Research Institute, Ayub Agricultural Research
Institute, Faisalabad, Pakistan for providing seed of sugarcane
genotypes.
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