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Documente Cultură
DOI: 10.1002/jpln.201400202
831
Review Article
School of Agriculture and Food Science/CRC CARE, The University of Queensland, St Lucia, Queensland 4072, Australia
Abstract
Our current understanding of silicon (Si) availability in agricultural soils is reviewed and knowledge gaps are highlighted. Silicon is a beneficial rather than essential plant nutrient and yield responses to its application have been frequently demonstrated in Si-accumulator crops such as
rice and sugarcane. These crops are typically grown on highly weathered (desilicated) soils
where soil solution Si concentrations are low. Increased yields are the result of simultaneous
increases in plant tolerance to a wide range of biotic (plant pathogens, insect pests) and
abiotic (water shortage, excess salts, metal toxicities) stresses. Traditionally, soil solution Si is
viewed as being supplied by dissolution of primary and secondary minerals and buffered by adsorption/desorption of silicate onto Al and Fe hydrous oxide surfaces. In recent years it has become recognized that phytogenic cycling of Si [uptake of Si by plants, formation of phytogenic
silica (SiO2 nH2O) mainly in leaves and subsequent return of this silica to soils in plant litter] is
the main determinant of soil solution Si concentrations in natural forests and grasslands. Considerable diminution of the phytogenic Si pool in agricultural soils is likely due to regular removal of
Si in harvested products. A range of extractants (unbuffered salts, acetate-based solutions, and
acids) can provide valuable information on the Si status of soils and the likelihood of a yield response in rice and sugarcane. The most common Si fertilizers used are industrial byproducts
(e.g., blast furnace slag, steel slag, ferromanganous slag, Ca slag). Since agriculture promotes
soil desilication and Si is presently being promoted as a broad spectrum plant prophylactic, the
future use of Si in agriculture is likely to increase.
Aspects that require future research include the role of specific adsorption of silicate onto hydrous oxides, the significance of phytogenic Si in agricultural soils, the extent of loss of phytogenic Si due to crop harvest, the role of hydroxyaluminosilicate formation in fertilized soils, and
the effect of soil pH on Si availability.
Key words: silicon / phytoliths / soil Si pools / soil desilication / fertilizer Si / biogenic Si
Introduction
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subsequent return to the soils in plant litter) drives soil solution Si concentrations in natural ecosystems; and this exportation of harvested product in agriculture severely depletes
soils of their phytogenic Si pool (Keller et al., 2012). Indeed,
because the continual removal of Si in harvested product is
promoting desilication of soils, addition of fertilizer Si may become a more common practice for a wide range of crops in
the future (Meena et al., 2014). In addition, because of its
known effect in protection of plants against biotic stresses, Si
is now being promoted and researched as a broad spectrum,
environmentally friendly plant prophylactic which can establish a broad spectrum resistance to diseases and pests in a
wide range of plants (Keeping and Kvedaras, 2008; Van
Bockhaven et al., 2013).
Little work has concentrated on the fertility aspects of Si availability in agricultural soils. Modern textbooks on soil fertility
normally mention Si in passing and there is no definitive coverage of its availability and crop use. The purpose of this paper is to provide a contemporary view of Si availability in agricultural soils and to point out where gaps in our knowledge
exist.
CaCl2-extractable Si / mg kg1
Soil pH
11
pH
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Plants differ greatly in their ability to accumulate Si and concentrations in above-ground parts can range from 1 to 100 g
kg1 (they are normally present in concentrations of similar order to those of macronutrients such as N and K) (Epstein,
1999; Ma and Takahashi, 2002). The process of Si uptake
can be classified as active, passive or rejective, and accordingly plants can be classified as accumulators, intermediates,
and excluders. Plants are considered as Si accumulators
when the tissue Si concentration is > 10 g kg1 and as excluders when they have concentrations < 5 g kg1 (Ma and
Takahashi, 2002). Seven (sugarcane, rice, wheat, barley, sugar beet, soybean,s and tomatoes) out of the 10 most important crops (ranked by global production) are classified as accumulators (Guntzer et al., 2012a).
The distribution of Si within shoot organs and tissue is determined primarily by the transpiration rate of the organ. Most of
the Si remains in the apoplast and is deposited at the termini
of the transpiration stream, mainly in the outer walls of the epidermal cells in the shoots. The deposited silicic acid is concentrated by transpiration loss of water and polymerizes as
amorphous silica (SiO2 nH2O) to form deposits known as
phytogenic Si (Smithson, 1956). These silica deposits can
contain small amounts of impurities such as Al, Fe, Ti, Mn, P,
Cu, N, and C (Clarke, 2003). Silica can also accumulate in
the vascular system and the endodermis of roots (Mitani and
Ma, 2005). Amorphous phytogenic Si is immobile within the
plant and so Si tends to accumulate in older plant tissues.
The application of Si often improves plant resistance to a
range of biotic and abiotic stresses (Epstein, 1999; Ma and
Yamaji, 2006; Liang et al., 2007; Meena et al., 2014). The
main biotic stresses alleviated include pathogen and insect
pest damage, while the most important abiotic stresses are
drought, salinity, and metal toxicities. Since plants growing in
the field are constantly subjected to an array of stresses, yield
increases due to fertilizer Si applications are not surprising on
Si-deficient soils. Silicon is typically applied as slag (see below) to rice at about 225300 kg Si ha1 annually and to sugarcane at about 6001125 kg Si ha1 every 45 y (at replanting) (Kingston, 2008). Yield responses under field conditions
to Si fertilizer additions in sugarcane have been reported as
1050% in Hawaii (Ayres, 1966; Clements, 1965; Fox et al.,
1967), 614% and 20% in Florida (Gascho and Andreis,
1974; Raid et al., 1992), 712% in Brazil (Korndorfer and
Lepsch, 2001), and 2141% in Australia (Kingston et al.,
2005). Similarly, for rice yield responses have been reported
as 14% and 1030% in Japan (Yoshida, 1981; Mizuochi,
2002), 0400% and 829% in China (Wang et al., 2001;
Wang, 2005), and 17% in South Korea (Ali et al., 2008). Yield
increases have also been observed in a wide range of grain
crops including wheat, barley, rye, and sesame, as well as
Importance of phytogenic Si
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Solubility/availability of phytogenic Si
1,E-12
Quartz
Amorphous silica
1,E-13
Soil phytoliths
Heated phytoliths
Fresh phytoliths
1,E-14
1,E-15
1,E-16
1,E-17
pH
Figure 3: Comparison of the effect of pH on the solubility of some typical soil clay minerals and parent rock primary minerals with that of
phytogenic Si. Redrawn from Fraysse et al. (2009).
10
12
14
pH
Figure 4: Comparison of the dissolution of phytogenic Si with that of
amorphous silica and quartz as a function of pH. Redrawn from
Fraysse et al. (2006).
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Estimation of plant-available Si
Table 1: Some selected methods used to determine extractable soil Si along with suggested critical levels below which a crop response can
be expected.
Extractant
H2 O
1:10 for 4 h
< 0.90
0.01 M CaCl2
1:10 for 16 h
< 20
1:50 for 1 h
< 20
1:10 for 1 h
< 15
Snyder (1991)
1:10 for 5 h
< 60
1:200 for 16 h
< 100
Hurney (1973);
Berthelsen et al. (2003)
< 4070
1:10 for 24 h
< 200300
(Table 1). They still extract more Si from soils with a higher Si
status and more from Si-fertilized rather than unfertilized soils.
Nevertheless, Berthelsen et al. (2001; 2005) suggested that
the best single measure of Si availability was that extracted
with 0.01 M CaCl2. In a survey of 200 sugarcane sites in N
Queensland, Berthelsen et al. (2003) found a close relationship between leaf Si and 0.01 M CaCl2-extractable Si, while
no relationship was obtained when 0.005 M H2SO4 was used
as the extractant. Similarly, in a survey of 28 sites in South
Africa, Miles et al. (2014) found a close correlation between
soil and leaf Si when 0.01M CaCl2 was used as an extractant
(R2 = 0.77) and a much weaker one (R2 = 0.48) when 0.02 M
H2SO4 was used.
Fertilizer Si
tive Si source for crops, diatomaceous earth has limited solubility in acidic conditions and its solubility increases greatly at
pH > 9.0 (as with phytogenic Si). Berthelsen et al. (2003)
showed that diatomaceous earth was generally an ineffective
source of Si for sugarcane grown in soils of pH 4.86.0.
The most common Si fertilizers are various industrial by-products. Useful materials from the steel industry include blast furnace slag, steel slag, converter slag, ferromanganous slag,
and silica fume. Calcium silicate slag is also produced as a
by-product of the manufacture of phosphorus in electric arc
furnaces. These slag materials contain various amounts of
Ca- and/or Mg-silicates and are very effective fertilizer materials as long as they do not contain significant amounts of
heavy metal contaminants (Gascho, 2001; Korndorfer and
Pereira, 2011). Slags also act as liming materials since dissolved OH ions are released. The dissolution of akermanite,
a major mineral component of blast furnace slag, is shown
below:
Ca2MgSi2O7 + 7 H2O > 2 H4SiO4 + Mg2+ + 6 OH + 2 Ca2+
Registration of Si slags as fertilizers in Japan requires the material to contain not less than 20% SiO2 and to have a neutralizing capacity of > 35%, 100% of particles < 200 mm and
> 60% of particles < 600 mm, and to conform to standards for
acid soluble and total loadings of Ni, Cr, and Ti (Takahashi,
2002; Owa, 2002). Standards developed in other countries
are similar (Kingston, 2008). A range of these slag materials
have been used very successfully as Si fertilizers for both rice
and sugarcane (Takahashi, 2002; Kingston, 2008).
There are also some manufactured fertilizer Si products including fused Mg-phosphate (7.59.4% acid soluble Si),
which is made by fusing a mixture of phosphate rock and olivine or serpentine, and fused K-silicate (1112% acid soluble
Si), which is made from calcining coal fly ash with K- and Mghydroxide. Silica gel is manufactured from Na-silicate and sulfuric acid, and its solubility can be increased by small particle
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Foliar applications
There has been commercial interest in developing siliconcontaining solutions (normally K or Na-metasilicate or silicic
acid) for foliar applications, particularly in relation to disease
and pest control (i.e., environmentally friendly fungicides and
pesticides). Their use has met with limited success. For example, there have been some reports of control of powdery
mildews on cucumber, muskmelon, zucchini, and grape
(Bowen et al., 1992; Menzies et al., 1992). As already noted,
plants (especially Si-accumulators) absorb Si in quantities
similar to those for macronutrients (e.g., N and K), and by
comparison with plant-requirement or the quantities that can
be soil-applied, the amounts that can be applied in foliar
sprays are very small. Thus, Matlou (2006) found that regular
foliar applications of Si to sorghum on a Si-deficient soil did
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Irrigation water
10 Conclusions
In the future, Si is likely to be applied to agricultural soils on a
much more extensive and frequent basis. This is because
continual removal of Si in harvested products will, over time,
decrease the Si status of many agricultural soils low in weath-
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Acknowledgments
I thank the late Professor K. M. Goh, to whom this paper is
dedicated, for stimulating my interest in the fertility of agricultural soils.
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