Journal of Plant Nutrition, 28: 917–929, 2005

Copyright © Taylor & Francis Inc.

ISSN: 0190-4167 print / 1532-4087 online
DOI: 10.1081/PLN-200058877

Effect of Silicon Application on Cadmium Uptake

and Distribution in Strawberry Plants Grown
on Contaminated Soils

Waldemar Treder1 and Grzegorz Cieslinski2

1
Research Institute of Pomology and Floriculture, Skierniewice, Poland
2
Agricultural University, Warsaw, Poland

ABSTRACT

Cadmium (Cd) is one of the most toxic metals affecting the environment. The initial
Cd concentration in soil could influence its uptake and distribution in plants. In this
paper, the effects of soil type and contamination level on Cd uptake and distribution
within the ‘Elsanta’ strawberry plant are presented. Silicon (Si) as potassium silicate,
applied by spraying or directly onto soil, was used to decrease Cd uptake and alleviate
it toxic effects. Plants were grown in a greenhouse in sandy and sandy clay-loam soils
contaminated with five different Cd levels. The concentrations of Cd in roots, stems,
leaves, and fruits were measured after fruit harvest. The results obtained showed that
the initial soil contamination and soil type significantly influenced Cd uptake and its
distribution within the strawberry plants. As expected, Cd uptake by strawberry plants
increased with the initial level of this element in the soil. However, plants grown on
sandy soil had a significantly higher concentration of Cd in all organs compared with
plans grown on sandy clay-loam soil. Silicon used as soil amendment prior to planting
was effective in preventing excessive Cd uptake by strawberry plants grown on sandy
soil. The lower Cd concentration as the effect of Si application was observed in stems,
leaves, and fruits but not in roots. A foliar application by spraying with potassium
silicate did not reduce the concentration of Cd in the individual parts of strawberry
plants.

Keywords: cadmium, Fragaria × ananassa Duch., nutrient concentration, heavy met-

als, soil pollution, silicon

Received 16 April 2003; accepted 10 August 2004.

Address correspondence to Waldemar Treder, Research Institute of Pomology and
Floriculture, 96-100 Skierniewice, Poland. E-mail: wtreder@insad.pl

917
918 W. Treder and G. Cieslinski

INTRODUCTION

The risk of food-chain contamination by toxic substances and elements has

been a major concern of both researchers and consumers. Heavy metals, es-
pecially cadmium (Cd), are one of these dangers (Galal-Gorchev, 1993; Kloke
et al., 1984; Prasad, 1995; Wagner, 1993). Cadmium is a natural element of
the lithosphere and its concentration in uncontaminated soils does not usu-
ally exceed 1 mg kg−1 of soil. Any positive effects of Cd on living organisms
have been proven so far, although it is very easily taken up by plant roots and
transported to the upper parts of plants (Kabata-Pendias and Pendias, 1992).
The concentration of Cd in plants reflects its concentration in the soil solution
(Braumemer et al., 1986; Gerritse et al., 1983; Sánchez et al., 1999; Barkelaar
and Hale, 2003; McBride, 2002; Huang et al., 2003). In most plant species,
Cd is accumulated in the roots, although the allocation to the plant organs may
vary considerably among different species. For example, in sugarbeet, 10%–
20% was transported to the shoot (Greger and Lindberg, 1986); in soybean,
only 2% of the accumulated Cd reached the leaves (Cataldo et al., 1981), but in
strawberry, 7% of total Cd taken up by the plant was found in the fruits (Treder
and Cieslinski, 2000). Strawberry plants are known for their enhanced, selective
Cd uptake and accumulation in the fruits (Treder and Cieslinski, 2000; Szteke
et al., 1989; Cieslinski et al., 1994). Excessive accumulation of this element
in fruits is observed even in soils that are naturally poor in Cd. The uptake of
Cd is also significantly dependent upon the plant’s genetic attributes, as it has
often been observed that various cultivars of the same species accumulated the
element in different concentrations (Florijn and Van Beusichem, 1993; Guo
and Marschner, 1996; Treder and Cieslinski, 2002). Thus, Cd concentration in
a plant and its distribution to various parts of the plant are the result of the
combined influence of soil properties and biological factors. However, translo-
cation of Cd from the soil solids to the soil solution, and thus its availability
to plants, depends upon several factors. Soil properties such as organic matter
content, clay type, redox status, and soil pH are considered the major factors
determining the bioavailability of heavy metals in soil (Sommers et al., 1987).
The role of the soil environment in the process of Cd uptake becomes even more
important when we consider how it may be changed relatively easily as a result
of natural biological and chemical phenomena taking place in the soil or by di-
rect human activity. Excessive selective uptake of Cd in strawberry plants leads
to its accumulation in fruits of levels dangerous for consumption by humans,
especially children. Therefore, there is a need to investigate the mechanism of
Cd accumulation in strawberry plants and to devise methods for minimizing
uptake of the element from the soil. Silicon (Si) is one of the most abundant
elements on the earth’s surface and is a common, but generally minor, element
in the majority of living organisms. The biological role of Si in higher plants
has not been clearly understood (Epstein, 1999). Silicon is readily taken up by
plants, in many cases against the concentration gradient (Barber and Shone,
 Effect of Si Application on Cd Uptake 919

1966; Jarvis, 1987) and actively transported to different plant organs (Rafi and
Epstein, 1999). Absorption and transport of Si could interfere with the uptake
of other elements. It has been reported that Si has some positive properties and
could be useful in detoxification of soils with excessive manganese (Mn) and
aluminum (Al) contents (Galvez et al., 1987; Corrales et al., 1997). Investiga-
tions of the uptake and accumulation of Cd in apple trees by Cieslinski et al.
(1997) indicated that the location and distribution of Cd in roots were the same
as those of silicon (Si). This suggests that Si may play an important role in Cd
metabolism in plants, especially in its metabolic inactivation. Silicon availabil-
ity to plants may influence the uptake and accumulation of Cd in strawberry
plants.
The purpose of this study was to examine the effect of Si application on
Cd uptake and concentration in strawberry plants grown on two different soil
types, contaminated at five different initial Cd levels.

MATERIALS AND METHODS

The experiment was carried out in the greenhouses of the Institute of Pomology
and Floriculture in Skierniewice (lat 51◦ 50 N, long 20◦ 20 E). The “frigo”
type strawberry plantlets cv. ‘Elsanta’ were planted in 0.9 L plastic pots. The
following two types of soil were used: sandy soil from Skierniewice and sandy
clay-loam soil from Research Station Brzezna, in south Poland. Some properties
of the soil are shown in Table 1. The soils were collected from the field to a
depth of 20 cm by hand using a shovel. The soils were allowed to air-dry before
being crushed to pass through 2 mm sieve, and then enriched with nitrogen (N),
phosphorus (P) and potassium (K) at 15, 8, and 35 mg kg−1 soil, respectively.
As a sources of N, P, and K, ammonium nitrate, monopotassium phosphate, and
potassium sulphate were used. In order to avoid introducing additional nutrients
or contaminants along with the fertilizers, only analytical-grade compounds
were used. Cadmium was added as CdSO4 8H2 0 to the air-dried soil prior to
planting at five application rates: 0, 4.4, 8.8, 13.2, and 17.6 µM Cd kg−1 of
soil. The soil samples with Cd were transferred to plastic pots and incubated in

Table 1
Properties of soils used in the experiment

Solid fraction
content (%)
Organic matter DTPA-extractable
Soil type Sand Silt Clay C (g 100 g−1 ) Cd (mg kg−1 ) pH (KCl)

Sandy 70.5 15.3 14.2 1.1 0.096 4.6

Sandy clay-loam 56.0 15.8 28.2 3.1 0.143 4.9
920 W. Treder and G. Cieslinski

a greenhouse (22◦ C day, 15◦ C night) for a period of four weeks to equilibrate
with the soil. The moisture level of the incubating soil was adjusted to 65%
of water capacity with distilled H2 O and maintained at this level throughout
the incubation. After the plants were individually planted into the pots, they
were watered with distilled water to keep soil water capacity at a level between
65% and 100%. The following treatments to the soil and plants of potassium
silicate were applied: C, control, without Si; S, 16.7 mM Si kg−1 of soil, applied
directly to the soil before planting; F, triple spray with a total rate of 16.7 mM
Si per plant (0.25% of potassium silicate); SF, silicon applied directly to the
soil at a rate of 16.7 mM Si per plant and triple foliar spray of 16.7 mM Si per
plant.
The experimental design was a split-plot block: 2 × 5 × 4. The treatments
consisted of: two soil types × five levels of soil Cd contamination and four
types of plant treatments with silicon. There were four replicates with eight
plants in each.
The fruit matured between the 11th and 14th week after planting. After
fruit harvest, the whole plants were lifted, washed with distilled water, sepa-
rated into different organs, and dried in an air-flow oven at 70◦ C. Fruit samples
were washed with distilled water and kept frozen until analysis. Cadmium con-
centration was measured separately in roots, crowns, leaves, and fruit. Dried
plant material was ground with a metal-free grinder and mineralized in con-
centrated nitric acid, whereas fruit samples were ashed in a muffle furnace at
550◦ C and the ash dissolved in hydrochloric acid. Determination of Cd in the
sample solution was made with an inductively-coupled plasma spectrometer
(IR-ICP, Thermo-Jarrel Ash, Franklin, Massachusetts, USA).
The data were processed by three-way ANOVA and the least significance
difference test (LSD) was used as a post-test. The data obtained were evaluated
using the statistical package STATISTICA 5,5 A (StatSoft, Inc., New York,
USA).

RESULTS AND DISCUSSION

The results of the present study showed that Cd concentration in different straw-
berry organs was influenced significantly by all investigated factors (Table 2).
The type of soil and soil contamination with Cd showed the significance of
the F rates for the concentration of Cd in the roots, stems, leaves, and fruits of
strawberry plants. Silicon application significantly decreased the concentration
of Cd in the stems, leaves, and fruits of strawberry plants, although it did not
influence the Cd concentration in the roots. The ANOVA analysis also showed
significant two-way and three-way interactions between investigated factors
on Cd concentration in individual parts of strawberries. However, in the case
of the roots and stems, we did not confirm the significance of the interaction
between Cd concentration in soil and Si application. Table 2 shows statistics
 Effect of Si Application on Cd Uptake 921

Table 2
Values of Femp. influence of investigated factors on Cd concentration in individual parts
of strawberry plants (cultivar “Elsanta”)

Source of variation Roots Stems Leaves Fruits

Type of soil (1) 32.48∗∗∗ 503.51∗∗∗ 332.20∗∗∗ 240.78∗∗∗

Cd concentration in soil (2) 84.19∗∗∗ 257.18∗∗∗ 162.50∗∗∗ 165.91∗∗∗
Si application (3) 1.55 NS 5.68∗∗ 7.24∗∗∗ 5.39∗∗
1×2 10.90∗∗∗ 62.60∗∗∗ 47.34∗∗∗ 26.25∗∗∗
1×3 2.78∗ 10.59∗∗∗ 9.59∗∗∗ 9.24∗∗∗
2×3 0.90 NS 1.82 NS 2.76∗∗ 2.38∗∗
1×2×3 2.30∗ 4.59∗∗∗ 5.23∗∗∗ 3.31∗∗∗
∗∗∗
Significant at 0.001; ∗∗ significant at 0.01; ∗ significant at 0.05; NS, non-significant
by F test.

confirming a highly significant soil effect on Cd concentration in strawberry

organs. This effect is also demonstrated by the interaction effects of soil type
× Cd concentration in the soil and soil type × Cd concentration in the soil ×
Si application.

Concentration of Cadmium in Roots

Initial contamination of both soil types with Cd significantly influenced its

concentration in the roots of strawberry plants (Table 3). Cadmium concentra-
tion in the roots varied widely, ranging from 2.2 mg kg−1 to almost 39.0 mg
kg−1 and increased with Cd level in the soil. However, plants grown on the
light, sandy soil had a higher concentration of Cd in roots than plants grown
on heavy, sandy-loam soil. This result was probably related to the lower

Table 3
Cadmium concentration in strawberry roots (mg kg−1 DW) depending on soil
contamination

Type of soil
Amount of Cd added to soil
[µM Cd kg−1 of soil] Sandy soil Sandy clay-loam soil Difference [d]

Control 2.22 4.01 1.79

4.4 11.95 8.02 3.93
8.8 21.41 15.60 5.81∗
13.2 38.97 18.05 20.92∗∗
17.6 32.83 28.80 4.03
∗
LSD (5 %) = 5.11; ∗∗ LSD (1 %) = 6.75.
922 W. Treder and G. Cieslinski

buffering capacity of sandy soils and thus higher availability of Cd to plants.

Plants on sandy soil showed the highest Cd concentration in roots (38.97 mg
kg−1 DW) when initial Cd was 13.2 µM Cd kg−1 , whereas plants on heavy soil
had the highest concentration (28.8 mg kg−1 D.W.) at the highest initial Cd soil
contamination—17.6 µM Cd kg−1 of soil. This difference could be explained
by higher toxicity of Cd due to its greater availability to the plants on sandy
soils. The capacity of roots to accumulate large amounts of Cd is well known
(Kabata-Pendias and Pendias, 1992; Gerritse et al., 1983; Treder and Cieslinski,
2000; Cieslinski et al., 1997). Cadmium concentration in the soil solution is the
main factor influencing the Cd uptake from soil. An increased Cd concentration
in soil resulted in its elevated level in roots (Braumemer et al., 1986). Silicon
application did not influence the Cd concentration in strawberry roots (data not
presented).

Concentration of Cadmium in Stems

Chemical analysis showed a significant differentiation of Cd concentration in

the stems of strawberry plants, which was related to all investigated factors.
Plants grown on the control—uncontaminated soil—had very low Cd concen-
tration in stems on both soil types (Table 4). Cadmium concentration increased
proportionally with increase of initial Cd contamination, but it was significantly
higher on sandy soil (Table 4). Cadmium concentration in stems on plants grown
on sandy soils was 4.1, 4.0, 5.0, and 2.6 times higher than on heavy, sandy-loam
soil with Cd doses of 4.4, 8.8, 13.2, and 17.6 µM kg−1 of soil, respectively.
The effects of Si application on Cd concentration in strawberry stems are not
very clear and generally depended on form of application (Figure 1). In the case
of sandy soil, there was a tendency for treatment SF (Si applied to soil before
planting and triple foliar spraying) to be effective, especially at the highest Cd
dose, 17.6 µM Cd kg−1 of soil. However, on heavy, sandy-loam soil, the effect

Table 4
Cadmium concentration in strawberry stems (mg kg−1 DW) depending on soil
contamination

Type of soil
Amount of Cd added to soil
[µM Cd kg−1 of soil] Sandy soil Sandy clay-loam soil Difference [d]

Control 0.79 0.84 0.05

4.4 11.76 2.81 8.95∗∗
8.8 17.74 4.46 13.28∗∗
13.2 33.83 6.81 27.02∗∗
17.6 47.77 18.42 29.35∗∗
∗
LSD (5%) =3.10; ∗∗ LSD (1 %) = 4.10.
 Effect of Si Application on Cd Uptake 923

Figure 1. Cadmium concentration in strawberry stems (mg kg−1 DW) depending on

soil contamination and Si application.

of this treatment was the opposite and caused the highest Cd concentration in
stem tissue. Such differentiation is typical for plant organs taking part in the
transport of minerals but not being their accumulators. In the case of straw-
berries, the stems, also called crowns, are very short. Thus, the concentration
of Cd in stems is not directly related to the accumulation ability of plant tis-
sues, as it shown by Cieslinski et al. (1997). The stems of strawberry plants
cultivated in uncontaminated soils usually had lower concentrations of Cd in
stems compared with roots. However, on the highest Cd dose on sandy soil, the
concentration of Cd in stems exceeded that in roots (Tables 3 and 4).

Concentration of Cadmium in Leaves

The Cd concentration in the strawberry leaves varied widely, ranging from 0.19
to 25.6 mg kg−1 DW, and it increased according to initial Cd contamination
in the soil (Table 5). Strawberry leaves generally had concentrations twice as
low as Cd in stems (Tables 4 and 5). As in the case of roots and stems, the Cd
concentration in leaves rose according to increasing Cd levels in soil and was
significantly higher on sandy soil. Cadmium concentration in leaves on plants
grown on sandy soils was 3.5, 3.5, 4.2, and 2.9 times higher than on heavy,
sandy-loam soil on initial Cd doses 4.4, 8.8, 13.2, and 17.6 µM kg−1 of soil,
respectively. The results of the present study confirm the earlier observation of
McLean (1976) that plants grown on soils contaminated with Cd and containing
relatively low levels of silt minerals and organic matter accumulated more Cd.
924 W. Treder and G. Cieslinski

Table 5
Cadmium concentration in strawberry leaves (mg kg−1 DW) depending on soil
contamination

Type of soil
Amount of Cd added to soil
[µM Cd kg−1 of soil] Sandy soil Sandy clay-loam soil Difference [d]

Control 0.19 0.57 0.38

4.4 6.08 1.73 4.35∗∗
8.8 9.93 2.83 7.10∗∗
13.2 19.80 4.62 15.18∗∗
17.6 25.60 8.83 16.77∗∗
∗
LSD (5%) =2.08; ∗∗ LSD (1 %) = 2.75.

It has been stated that not only the silt fraction but also organic substances
in soil may restrict the availability of Cd to plants (McLean, 1976). Thus,
cultivation of strawberry plants in sandy, poor soils that are contaminated with
Cd may consequently lead to hyper-accumulation of Cd in plants as well as
in fruits (Treder and Cieslinski, 2000; Cieslinski et al., 1994; Szteke et al.,
1989; Cieslinski et al., 1996). Silicon in the form of foliar spraying and soil
application (SF) as potassium silicate significantly influenced the decrease in
the concentration of Cd only in the leaves of strawberry plants cultivated on
highly contaminated sandy soil—13.2 and 17.7 µM Cd kg−1 (Figure 2). At the

Figure 2. Cadmium concentration in strawberry leaves (mg kg−1 DW) depending on

soil contamination and Si application.
 Effect of Si Application on Cd Uptake 925

highest Cd levels in sandy soil (17.7 µM Cd kg−1 ), all kinds of treatment with
Si were effective in lowering Cd levels in leaves as compared with the control
(non-treated plants). A positive influence of Si on plants has been reported in
many papers concerning the role of this element in enhancing the tolerance of
plants to pathogenic fungi and pests (Kunoh et al., 1975; Carver et al., 1987).
The results of earlier studies also showed that the increased availability of Si
to plants in the soil solution had improved the supply of P to plants (Miyake
and Takahashi, 1978) and diminished the phytotoxicity of Mn and Al (Galvez
et al., 1987; Corrales et al., 1997).

Concentration of Cadmium in Fruits

Cadmium application before planting significantly increased its concentration

in the fruits of strawberry plants on both soil types. The concentration of Cd in
strawberry fruits varied from 0.015 on both soil types for the control treatment,
without Cd application, to 0.958 mg kg−1 of FW on sandy soil at the highest
initial Cd dose, 17.6 µM Cd kg−1 soil (Table 6). However, Cd concentration in
fruits on plants grown on sandy soils was 4.7, 3.5, 3.5, and 2.3 times higher than
on heavy, sandy-loam soil with initial Cd doses of 4.4, 8.8, 13.2, and 17.6 µM
kg−1 of soil, respectively. As in the case of other strawberry organs, comparing
the differences between Cd accumulations on both soil types, it decreased with
increasing Cd contamination in soil. This finding means that at lower doses of
Cd, higher buffering capacity of heavy soil prevented excessive Cd uptake by
plants. However, with increasing Cd level up to 17.6 µM Cd kg−1 of soil, the
potential ability of heavy soil to absorb Cd decreased and thus allowed higher
Cd uptake by plants. There was a significant interaction among the type of soil,
its contamination with Cd, and Si application. Silicon application directly to
soil (S) and foliar and soil treatment (SF) in the form of potassium silicate,

Table 6
Cadmium concentration in strawberry fruits (mg kg−1 FW) depending on soil
contamination

Type of soil
Amount of Cd added to soil
[µM Cd kg−1 of soil] Sandy soil Sandy clay-loam soil Difference [d]

Control 0.015 0.015 0.00

4.4 0.258 0.055 0.203∗∗
8.8 0.347 0.099 0.248∗∗
13.2 0.548 0.157 0.391∗∗
17.6 0.958 0.418 0.540∗∗
∗
LSD (5%) =0.079; ∗∗ LSD (1 %) =0.104.
926 W. Treder and G. Cieslinski

Figure 3. Cadmium concentration in strawberry fruits (mg kg−1 FW) depending on soil
contamination and Si application.

significantly decreased Cd concentration in fruits, especially on sandy soil at

doses of 4.4, 8.8, and 13.2 µM Cd kg−1 of soil (Figure 3). At the highest
Cd contamination level, 17.6 µM Cd kg−1 of soil, only the treatment with Si
applied to soil prior to planting (S) was efficient (Figure 3). Individual foliar
spraying with potassium silicate did not provide such an effect. Taking into
consideration the fact that Si in this experiment caused a significant decrease
in the concentration of Cd in the leaves and fruits of strawberry plants and
the fact that it is taken up and transported along with water and then stored in
cell walls, where it enhances their tightness and stiffness, it seems likely that
Si is a natural mechanical obstacle for the ions of Cd in their inner ways of
transport to cells. Such a mechanism has been observed by Wang et al. (2000)
in their investigation of Cd uptake by rice plants. In this paper, the authors
showed that the mechanism of detoxification consists of precipitation of Si and
Cd complexes in the cell walls and in the intercellular spaces, which greatly
restricts the translocation of Cd from the roots to distant plant organs. Other Si
compounds, also deposited in cell walls, may restrict the transport of Cd in plant
tissues. A similar mechanism was observed using EDX (Energy-dispersive X-
ray microanalysis) in the restriction of availability of healthy plant tissue to the
mycelium of Erysiphe graminis in barley (Carver et al., 1987).
The results obtained showed the possibility of using Si as an efficient fac-
tor limiting Cd uptake by strawberry plants grown on sandy soils. It seems
that applying Si directly to soil, which can be done using fertilizers contained
Si, could be an effective way of preventing excessive Cd uptake during in-
tensive strawberry production on light soils. Due to the increasing popularity
 Effect of Si Application on Cd Uptake 927

of strawberry production using plastic bags and containers in greenhouses to

extend the production season, the quality of soil used for this purpose and es-
pecially its heavy-metals content should always be evaluated. The possibility
of using Si as a soil amendment before planting could be an effective way to
prevent excessive Cd uptake; however, this subject needs further research.

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