A Contemporary Overview of Silicon Availability in Agricultural Soils

J. Plant Nutr. Soil Sci.
2014, 177, 831844
DOI: 10.1002/jpln.201400202
831
Review Article
A contemporary overview of silicon availability in agricultural soils

Richard J. Haynes1*
1
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
Accepted September 19, 2014
Introduction
Silicon (Si) is the second most abundant element of the

earths crust with a mean content of 28.8% (Wedepohl,
1995). It occurs in more than 370 rock-forming minerals and
is a basic component of most soils. In recent times the global
Si cycle has received increased attention because of its interrelationship with the C cycle. Indeed, biogeochemical cycling
of Si regulates atmospheric CO2 concentrations via several
mechanisms including (1) chemical weathering of minerals in
the terrestrial environment, with consumption of CO2, and the
eventual precipitation of CaCO3 as marine sediments, (2) occlusion of C in phytogenic silica deposited in soil, and (3) biogenic sequestration of C in oceanic phytoplankton (Si is an
essential nutrient for growth of diatoms) and their subsequent
sedimentation (Street-Perrott and Barker, 2008; Struyf et al.,
2009; Carey and Fulweiler, 2012; Frings et al., 2014). Of late,
the importance of Si to plant ecology in terrestrial systems
has been recognized (Cooke and Leishman, 2011) and it has
been suggested that removal of Si in harvested products in
agricultural systems is creating a new removal mechanism for

Si which is substantially changing the terrestrial and, thus,
global Si cycle (Vandevenne et al., 2012). Indeed, from the
viewpoint of managing agricultural ecosystems, Si is an agronomically important nutrient.
Although Si is still not considered as an essential nutrient, its
beneficial effects on plant growth and yields have been repeatedly demonstrated particularly for crops such as sugarcane and rice (Savante et al., 1997; 1999). Positive yield responses occur particularly in soils containing low concentrations of Si in soil solution including highly weathered Oxisols
and Ultisols and organic soils (e.g., Histisols). The main
mechanisms of yield increase are through improving plant resistance against abiotic (metal toxicities, water stress, salinity) and biotic (plant pathogens and insect pests) stresses
(Ma and Yamaji, 2006; Meena et al., 2014; Tripathi et al.,
2014). The Si status of agricultural soils has come under recent scrutiny due to the realization that phytogenic cycling of
Si (through the formation of phytogenic silica in plants and its
* Correspondence: Prof. R. J. Haynes; e-mail: r.haynes1@uq.edu.au
2014 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim
www.plant-soil.com
832
Haynes
J. Plant Nutr. Soil Sci. 2014, 177, 831844
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.
Silicon status of soils
A major consequence of the chemical weathering of primary

silicate minerals is desilication. Weathering releases highly
mobile basic cations (Ca2+, Mg2+, K+, and Na+), moderately
mobile Si(OH)4, and relatively immobile Al and Fe into soil
solution (White, 1995; Karathanasis, 2006). Part of the Si
released from the mineral structure reacts with Al (and to a
lesser extent Fe and Mg) to form secondary clay minerals,
while the remainder is subjected to leaching. As a consequence, most soils experience a loss of Si and basic cations
during weathering (White, 1995). Desilication is most pronounced in humid tropical environments and occurs to a lesser extent in temperate regions (Karathanasis, 2006). From
the viewpoint of this review, it is important to note that rice
and sugarcane are tropical/subtropical crops and the majority
of their production is on highly weathered Ultisols and Oxisols

(Savante et al., 1997; 1999), which have a characteristically
low Si status.
The main determinant of concentrations of Si in soil solution
is the solubility of primary and secondary minerals (Karathanasis, 2002; Sommer et al., 2006). Silicon in soil solution is
mainly present as monomeric silicic acid (H4SiO4) (Drees et
al., 1989; Dove, 1995; Dietzel, 2000) although at high pH
(> 9) silicic acid dissociates to form H3SiO4 and above pH 11
to H2SiO24 . Silicon concentrations in soil solution have been
reported to range from 0.01 to 2.0 mM (Karathanasis, 2002)
but are commonly in the range of 0.1 to 0.6 mM (Epstein,
2001; Sommer et al., 2006). In general, Si concentration in
soil solution decreases with increasing soil development
(Karathanasis, 2002; Sommer et al., 2006). As a result, the
soluble Si content of tropical soils, such as Ultisols and Oxisols, is generally 510 times less than that of most temperate
soils (McKeague and Cline, 1963; Juo and Sanchez, 1986;
Foy, 1992). Where a large group of soils in a region of a country is surveyed, there is generally a positive relationship between pH and Si solubility/extractability (Fox et al., 1967;
Cheong et al., 1968; Oya and Kina, 1989; Oya et al., 1989;
Miles et al., 2014). For example, the positive relationship between pH and CaCl2-extractable Si for 112 sites on the South
African sugar belt is shown in Fig. 1. The more weathered
soils with lower soluble Si are also those with lower pH values.
Silicate can be adsorbed to the surfaces of variably charged
soil colloids such as Fe and Al hydrous oxides. Such adsorption is highly pH-dependent because with increasing pH (up
to pH 9.8; the pK1 for silicic acid) an increasing proportion of
total Si in solution is present as H3SiO4 rather than H4SiO4.
Because Si is preferentially adsorbed in the anionic H3SiO4 form, maximum Si adsorption occurs at a soil pH of about 9.8
(Bowden et al., 1980). The relationship between pH and ionic
strength on the adsorption of silicate by the iron oxide goethite is shown in Fig. 2. As a result, in short-term equilibrium
Silicate adsorbed / mol cm2
CaCl2-extractable Si / mg kg1
Soil pH
Figure 1: Relationship between pH and CaCl2-extractable Si for 112

soils collected from sugarcane production fields in South Africa.
Redrawn from Miles et al. (2014).
11
pH
Figure 2: Effect of pH and ionic strength on adsorption of silicate by

goethite. Redrawn from Bowden et al. (1980).
www.plant-soil.com

experiments Si solubility in soils decreases with increasing
pH (Hingston and Raupach, 1967; Hingston et al, 1972;
Goldberg and Glaubig, 1988; Haynes et al., 2013). Thus, if
adsorption reactions are the major factor determining soil solution Si concentrations then Si solubility and availability will
decrease with increasing pH.
Silicon uptake and use by crops
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
Silicon availability in agricultural soils 833

horticultural crops such as garlic, onion, cucumber, chinese
cabbage, strawberries, and potatoes (Epstein, 1999; Voogt
and Sonnenveld, 2001; Kang and Jung, 2002).
Silicon has been shown to protect plants against a suite of
pathogens (mainly fungi but also some bacteria) (Currie and
Perry, 2007; Guntzer et al., 2012a; Van Bockhaven et al.,
2013). The increased resistance to disease occurs through
two main mechanisms. Firstly, impregnation of epidermal cell
walls with silica creates a hard outer barrier which mechanically impedes penetration by fungi and thereby disrupts the
infection process. Secondly, soluble Si can act as a modulator
of host resistance to pathogens through physiological and
biochemical/molecular mechanisms (Fauteux et al., 2006;
Brunings et al., 2009; Ghareeb et al., 2011; Van Bockhaven
et al., 2013). Similarly, Si applications can enhance plant resistance to insect pests (Laing et al., 2006; Kvedaras and
Keeping, 2007; Keeping and Kvedaras, 2008; Kvedaras et
al., 2010). The deposition of silica in epidermal layers offers a
physical barrier to sucking insects and leaf-eating caterpillars
and, in addition, Si plays a role in the induction of chemical
defences against herbivores (Correa et al., 2005; Gomes et
al., 2005; Ye et al., 2013).
It has been widely reported that the addition of Si can increase drought tolerance in crops including rice, sorghum,
wheat, sunflower, cucumber, and pepper (Liang et al., 2007;
Sacala 2009; Tripathi et al., 2014; Zhu and Gong, 2014). Silicon additions have also been shown to increase salinity tolerance in a similar range of crops (Meena et al., 2014; Tripathi
et al., 2014; Zhu and Gong, 2014). Such Si-stimulated tolerance to drought/salinity is achieved by a range of physiological (e.g., modulation of transpiration rates, increased photosynthesis, decreased uptake of Na+ and Cl) and biochemical/molecular (e.g., improved antioxidant defences, osmotic
adjustment with organic solutes) mechanisms (Zhu and
Gong, 2014).
Silicon applications can be effective at alleviating toxicity of
metals including Al, Mn, Fe, Cd, Pb, and Zn (Gu et al., 2012;
Maksimovic et al., 2012; Bharwana et al., 2013; Dufey et al.,
2013; Nhan and Hai, 2013). The responsible mechanisms are
complex and can be divided into external (e.g., co-deposition
of metals and Si in soil solution) and internal (e.g., co-deposition of metals and Si in roots, inhibition of root-to-shoot transport of metals) (Liang et al., 2007).
Importance of phytogenic Si
As already noted, Si accumulates in plants as silica deposits

particularly in the shoots. This silica is returned to the soil
mainly in plant litter, and as the organic matter (OM) is decomposed the silica is released. Extraction and analysis of
phytogenic Si in soils is often restricted to silt-sized particles
(this fraction is classified as phytolith Si) and the < 5 mm fraction is not included (Sommer et al., 2006). The term phytogenic Si is used here to describe all of the silica returned in plant
material. Due to the recycling of Si in the litter, the distribution
of phyogenic silica through the soil profile is typically similar
to that of organic C. Under natural forest or grassland there is
a pronounced accumulation at the soil surface and a rapid dewww.plant-soil.com
834
Haynes
crease with depth (Alexandre et al., 1997; Saccone et al.,

2007; 2008), while under arable cultivation (with regular inversion of the plough layer) both OM and phytogenic Si are more
evenly distributed within the surface layer (Guntzer et al.,
2012b; Seyfferth et al., 2013).
In recent years, the importance of cycling of Si through the
phytogenic Si pool in the soil of natural ecosystems has become recognized (Cornelis et al., 2011; Sommer et al., 2013;
Frings et al., 2014). Indeed, in natural forests and grasslands
nutrients are characteristically cycled from soil to plant and
then back to the soil, and losses are relatively small compared to the quantities continually being cycled. The flux of Si
through the plant in temperate and tropical forests ranges
from 576 (often 2050) kg ha1 y1 (Bartoli, 1983; Lucas et
al., 1993; Alexandre et al., 1997; Cornelis et al., 2010;
Meunier et al., 2010) and in grasslands it is in the same order,
12127 kg ha1 y1 (Blecker et al., 2006; Alexandre et al.,
2011; White et al., 2012).
Studies in various temperate and boreal forests have revealed that the immediate source of silicic acid taken up by the
trees is predominantly phytogenic silica and that this is then
converted back to phytogenic Si in tree foliage and then returned to the soil as tree litter (Farmer et al., 2005). Alexandre
et al. (1997), using a steady-state approach, showed that
about 74% of soil solution Si under a rainforest in the Congo
originated from phytogenic Si, while Bartoli (1983) found 85%
of Si uptake by a deciduous forest originated from dissolution
of phytogenic Si. Alexandre et al. (1997) calculated that about
92% of phytogenic Si is rapidly recycled, while only about 8%
are sequestered in a stable phytogenic pool. Gerard et al.
(2008) found 60% of the biogeochemical cycle of Si in a
Douglas fir forest originated from phytogenic Si. However, as
noted by Cornelis et al. (2011), the importance of the phytogenic cycling of Si will be highly dependent on the extent to
which mineral weathering has taken place. In a humid tropical
climate, in which there has been strong depletion of litho-pedogenic Si, phytogenic cycling of Si will supply the bulk of soil
solution Si and therefore that taken up by the plants.
The quantities of phytogenic Si measured in the soil varies
greatly between studies and may well be affected by the
method of measurement and soil type, as well as vegetation
cover (Keller et al., 2012). The range for natural forests (generally reported to a soil depth of 3050 cm) is 7,00031,262
(often 1,00018,000) kg Si ha1 (Alexandre et al., 1997;
Gerard et al., 2008; Saccone et al., 2008; Clymans et al.,
2011) and that for native grasslands 18,00071,100 kg Si
ha1 (Blecker et al., 2006). The range for the small number of
studies on agricultural land is 80013,458 kg Si ha1 (Desplanques et al., 2006; Clymans et al., 2011; Guntzer et al.,
2012b). Some diminution of phytogenic Si is to be expected
in agricultural soils (Keller et al., 2012; Struyf et al., 2010; Vandevenne et al., 2012). That is when a soil is converted to agriculturethe phytogenic Si pool (accumulated under natural
vegetation) is in disequilibrium with the current vegetation.
For example, after deforestation, Struyf et al. (2010) concluded that there are likely to be increased Si fluxes, depletion
of phytogenic Si, and eventually reduced Si fluxes. Similarly,
Sommer et al. (2013) observed considerable dissolution of

the phytogenic Si pool in forest soils due to disequilibrium
caused by loss of Si-rich grasses during canopy closure and
logging of trees. For agriculture, disequilibrium is mainly attributable to the high uptake of Si by some crops and the annual removal of much of this Si in the harvested product. Indeed, Matichenkov and Bocharnikova (2001) calculated that
210224 m tonnes of Si are removed from agricultural soils
each year.
The uptake by rice and sugarcane of 200400 kg Si ha1 y1
is well above that normally absorbed in natural systems (e.g.,
20100 kg Si ha1 y1) and, furthermore, the bulk of this phytogenic Si is removed from the system at harvest rather than
being returned to the soil. In S and SE Asia, where most of
the rice is grown, rice straw and husks are typically removed
from the field and used for a range of purposes, including animal fodder and fuel, so that all the above-ground vegetation
(and phytogenic Si) is removed from the field. This depletes
the soil of Si and accelerates soil desilication (Seyfferth et al.,
2013). For rice, retention of straw on the field (rather than
straw removal) is an important management tool for recycling
Si. This is because a large proportion of the above-ground Si
uptake (about 80%) is retained in the straw rather than the
harvested grain. Dobermann and Fairhurst (2000) calculated
that 4070 kg ha1 Si was held in 1 Mg of rice straw. For sugarcane, return of bagasse (the crushed cane) or bagasse fly
ash (if bagasse is used as a fuel source at the mill) and filter
mud to the fields can be important in maintaining the Si status
of soils (Savante et al., 1999).
Several workers have recently attempted to quantify the loss
of phytogenic Si from soils under agriculture. Clymans et al.
(2011) compared phytogenic Si in soils under boreal forest
with those under agriculture and measured larger quantities
under forest. They attributed the depletion under agriculture
to systematic removal of crop residues (and therefore a reduced return of phytogenic Si). Similar findings were reported
by Struyf et al. (2010) for temperate forest soils converted to
agriculture. On the classical Broadbalk continuous wheat experiment at Rothamsted, Guntzer et al. (2012b) measured a
decrease in phytogenic Si in the surface 23 cm of soil of between 19 and 66% between 1893 and 2000 (depending on
method of estimation). Keller et al. (2012) found that on a
long-term trial in Boigneville, France, a period of 12 y of wheat
straw removal (compared to straw returned) resulted in a reduction of 4164% in phytogenic Si in the surface 20 cm of
soil. Keller et al. (2012) calculated that, assuming that phytogenic Si was the only source of Si for crops and with an exportation of 50100 kg Si ha1 y1 and an initial pool of phytogenic Si of 1 Mg ha1, under a temperate climate and an initial
forest setting the phytogenic Si pool would be depleted within
a few decades. Desplanques et al. (2006) measured an exportation of Si from a rice field in the Camargue, France, by
grain and straw as 270 kg ha1 y1 and calculated that if phytogenic Si was considered the only source of Si for plants
then the stock of available Si would be exhausted after only
5 y. However, there were significant inputs of Si by irrigation
water. Yet, no workers have attempted to quantify the effects
of sugarcane monoculture on the phytogenic Si content of
soils. Nonetheless, Berthelsen (1997) compared a site under
rainforest with an adjacent long-term (30 y) sugarcane site in
www.plant-soil.com

N Queensland and found CaCl2-extractable Si levels were
13.1 mg kg1 under forest and only 5.3 mg kg1 under sugarcane. Such results demonstrate that under sugarcane there
was a decrease in Si supplying ability of the soil, which could
well be due to a decrease in phytogenic Si.
Solubility/availability of phytogenic Si
As discussed above, phytogenic Si is now recognized as one

of the most important sources of Si in soil solution among all
soil minerals (Fraysse et al., 2006; 2009; 2010; Nguyen et al.,
2014; Seyfferth et al., 2013; Gocke et al., 2013). It is, however, important to note here that phytoliths can be highly resistant to dissolution and may remain in soils for thousands of
years, thus, contributing to C sequestration (Parr and Sullivan, 2005) and their use in paleobotanical and archaeological
studies (Rovner, 1971; Santos et al., 2010). Fraysse et al.
(2009) compared the Si release rate from phytogenic Si to
that of typical primary [olivine, diopside (pyroxene), albite-orthoclase (feldspar)] and secondary (e.g., illite, kaolinite and
smectite) silicate minerals and found that at common soil pH
values (i.e., 48) phytogenic Si was 102 to 104 times more
soluble than the other minerals (Fig. 3). While the dissolution
rate of litho-pedogenic Si, in the form of primary and secondary minerals, decreases markedly with increasing pH (and is
greatly favored by acidic soil conditions), the reverse is the
case for phytogenic Si (Figs. 3 and 4). Dissolution of phytogenic Si increases with decreasing particle size due to the increase in surface area (Drees et al., 1989) and is promoted
by moist soil conditions and removal of soil solution Si.
Dissolution of phytogenic Si, measured between pH 2 and
12, was found by Fraysse et al. (2006) to be intermediate between that of quartz and of amorphous silica (Fig. 4). Dissolution rates of phytogenic Si originating from different plants
(larch, elm, fern, horsetail, and bamboo) over the same pH
range were very similar (Fraysse et al., 2006; 2009). Dissolution of SiO2 is due to nucleophilic attack of water molecules

that cause the breaking of siloxane bonds, >SiOSi<, at the
particle surface (Dove et al., 2008). Increasing pH leads to
the deprotonation of surface silanol groups, thereby facilitating the further breaking of the bridging siloxane bonds. Basepromoting dissolution occurs when the solution pH exceeds
the point of zero charge (for SiO2 in the pH range 1.24.0)
(Fraysse et al., 2006). Fraysse et al. (2009) calculated the
half-life of phytogenic Si in soil, assuming 100% water saturation over the year, to be 26156 weeks at pH 45 and only
710 weeks at pH 67. Similarly, Nguyen et al. (2014)
showed that Si release from phytogenic Si in rice straw was
an order of magnitude greater at pH 6.5 than at 3.0.
The sorption of di- and trivalent metals (Al3+, Fe3+, Zn2+,
Cu2+) to the surfaces of phytogenic Si is known to markedly
reduce its solubility (McKeague and Cline, 1963; Wilding et
al., 1979; Dove, 1995; Van Bennekom et al., 1991), and Al3+
has been shown to be particularly effective (Nguyen et al.,
2014). The cations are adsorbed on deprotonated Si-O sites
and mitigate water attack on SiOSi bonds. Fe3+ has a similar effect to Al3+ and a lowered decomposition rate of phytogenic Si in soils over time has been attributed to adsorption of
Al3+ and Fe3+ to its surfaces (Wickramasinghe and Rowell,
2006; Nguyen et al., 2014). Since the solubility of Al and Fe
increase greatly with decreasing pH (particularly below pH
5.5), their adsorption to the surfaces of phytogenic Si will be
favored at low soil pH. This will further reduce the solubility of
phytogenic Si at low soil pH and reinforce the trend for its increasing solubility with increasing pH. The adsorption of Al
and Fe to phytolith surfaces may partly explain why some can
be so persistent in soils.
Under field conditions, high rainfall conditions (as occur in
tropical regions where crops such as sugarcane and rice are
grown) will favor dissolution of phytogenic Si and dissolution
will occur most rapidly from smaller silica particles. A logical
extension of the above discussion is that liming will tend to in-
1,E-12
Quartz
Log R / mol Si g1 day1
R / mol Si cm2 sec1
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).
www.plant-soil.com
836
Haynes
crease the solubilization and plant-availability of phytogenic

Si. Yet, no such relationship between liming, phytogenic Si
solubility and plant availability of Si has been demonstrated.
Indeed, an increasing pH will also favor an increased adsorption of silicate by Al and Fe hydrous oxide surfaces and decreased weathering of primary and secondary minerals so
the overall effect is unclear. Further work in this area is warranted.
dispersion of soil particles (Berthelsen and Korndorfer, 2005)

meaning that the extracted solution may require filtering
through a micropore filter (after centrifugation) prior to analysis. Unbuffered salt solutions such as 0.01 M CaCl2 are favored (Berthelsen et al., 2003; Hohn et al., 2008; Miles et al.,
2014) since they have an ionic strength similar to that of the
soil solution and the dominant cation is Ca2+, as is the case in
most soil solutions. Their ionic strength also prevents dispersion and facilitates easy extraction and analysis.
Ammonium- and Na-acetates (buffered at low pH with acetic

acid) as well as acetic acid itself have been extensively used
as extractants for soil Si (Sauer et al., 2006). The acid extraction would result in dissolution of amorphous Al and Fe oxides
and hydroxides with release of adsorbed Si (Sauer et al.,
2006; Hohn et al., 2008) plus dissolution of amorphous aluminosilicates and any highly soluble crystalline aluminosilicate
material, while the presence of the acetate anion might favor
desorption of adsorbed silicate. In some cases phosphate
has been added to the Na-acetate extractant in order to displace more adsorbed silicate from adsorption surfaces (Fox
et al., 1967). In this regard, Heinai and Saigusa (2006) successfully used a phosphate buffer extraction (Na2HPO4) to
displace adsorbed Si for soil testing in paddy rice.
Estimation of plant-available Si
There are three key parameters that are likely to be important

in relation to the plant- availability of Si. These are (1) the intensity (concentration of Si in soil solution), (2) the capacity
(the reserve supply in solid phase that supplies the soil solution), and (3) the retention capability of the soil (Si adsorption
capacity). For example, for soil P a soil test extractant is generally used to measure a combination of intensity and capacity and, thus, whether P fertilizer is required or not, while a
P retention test is used to modify the fertilizer recommendation (the greater the P retention the more P needs to be
added) (Haynes, 1984). However, for Si the relative importance of various soil pools in relation to capacity is much less
clear than for P and probably differs in different situations
(e.g., dissolution of amorphous silicates, crystalline phyllosilicates, phytogenic Si or residual fertilizer Si), while the role of
Si adsorption is also unclear. For example, its adsorption
maximum is at pH 9.8, yet most soils, in which Si is deficient,
are in the pH range of 56. In contrast to P, as phosphate is
held by soil colloids so strongly that it is highly immobile in
soils, leaching of silicic acid from soils is a common phenomenon (Sommer et al., 2006; Gerard et al., 2008; Frings et al.,
2014). Although Gallez et al. (1977) developed a method for
measuring Si retention by soils (at pH 9.2) and Herbillon et al.
(1977) developed an index of Si saturation based on both
the Si adsorption and Si release characteristics, these parameters have not met with wide acceptance. In acid soils, precipitation of added Si in form of hydroxyaluminosilicates (see below) might well be a more important sink for Si than adsorption to soil colloids. There is a need for more detailed studies
into the chemistry of Si in soils before factors and interactions
influencing Si availability are fully understood.
Given the above uncertainties, it is not surprising that there is
no universally accepted method to evaluate plant-available Si
in soils. Information summarized in Table 1 illustrate the diversity of extractants that have been used. These tests have
been developed by groups of workers in different countries
and regions to meet the specific needs of the climates, soils,
and crops (mainly sugarcane or rice) in their locality. Many
other extractants have also been used although critical values
have not necessarily yet been derived (Berthelsen and Korndorfer, 2005; Sauer et al., 2006).
Water has been used by some workers to estimate readilysoluble Si (Fox et al., 1967; Elawad et al., 1982). However,
extracting with water dilutes the ionic strength of the soil solution greatly so that the amounts extracted will differ from those
present in soil solution. For example, as shown in Fig. 2, Si
adsorption onto metal hydrous oxides increases with increasing ionic strength. In addition, the low ionic strength results in
Citric acid has been suggested by some workers (Acquaye

and Tinsley, 1965), and the modes of extraction are likely to
include solubilization of Al and Fe hydrous oxides and aluminosilicates, displacement of adsorbed silicate by citrate and
complexation of Fe and Al by citrate, thus, preventing formation of Al and Fe complexes with Si during extraction (Sauer
et al., 2006). In addition, sulfuric acid has been used by a
number of workers (Hurney, 1973; Fox et al., 1967; Meyer
and Keeping, 2001). This is a modification of the Truog extractant for soil P, and in some cases both available P and Si
are determined on the same extract.
In general, acids such as sulfuric and citric acid extract more
Si than the Na- and NH4-acetate-based extractants, which in
turn extract more Si than water and CaCl2 (Fox et al., 1967;
Berthelsen et al., 2003; Barbosa Filho et al., 2004; Wang et
al., 2004; Kanamugire et al., 2006; Haynes et al., 2013). It
has been pointed out (Haynes et al., 2013) that acid extractants can remove very large amounts of Si from soils treated
with slags since these materials are acid-soluble. Some of
the Si extracted originates from residual unreacted slag
present in the soil. Similar observations have been made regarding the Na-acetate buffer (pH 4.0) method (Imaizumi and
Yoshida, 1958) by both Sumida (2002) and Wang et al.
(2001). These strongly acidic extractants should therefore be
avoided on Si-fertilized soils.
Many different extractants can provide valuable information
as to the Si status of soils and the likelihood of a yield response in rice or sugarcane. Indeed, when a range of different extractants are compared over a range of soils and fertilizer histories there is frequently a very strong correlation between extractable Si values derived from different extractants
(Berthelsen, 2000; Berthelsen et al., 2003, Wang et al., 2004;
Kanamugire, 2007). This occurs even though extractants
may remove widely different quantities of Si from the soil
www.plant-soil.com
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
Extraction ratio and period
Suggested critical level Reference
H2 O
1:10 for 4 h
< 0.90
Fox et al. (1967);

Elawad et al. (1982)
0.01 M CaCl2
1:10 for 16 h
< 20
Haysom and Chapman

(1975); Berthelsen et al.
(2003)
0.5 M NH4-acetate (pH 4.8)
1:50 for 1 h
< 20
Fox et al. (1967)
0.5 M acetic acid
1:10 for 1 h
< 15
Snyder (1991)
0.18 Na-acetate (plus 0.87 M acetic acid, pH 4.0)
1:10 for 5 h
< 60
Imaizumi and Yoshida (1958)
0.005 M sulfuric acid
1:200 for 16 h
< 100
Hurney (1973);
Berthelsen et al. (2003)
0.025 M sulfuric acid
1:10 for 20 min
< 4070
Meyer and Keeping (2001);

Kanamugire (2007)
0.04 M Na2HPO4 (pH 6.2)
1:10 for 24 h
< 200300
Heinai and Saigusa (2006)
(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
There is a range of silicate-containing materials suitable for

use as fertilizers (Gascho, 2001). In general, their usefulness
is dependent on reactivity rather than total Si content. There
are a few naturally occurring mineral materials such as wollastonite (2125% Si; CaSiO3), olivine (ca. 20% Si; MgSiO3)
and diatomaceous earth (3842% Si) that can be mined from
the earths surface and used. Availability often limits their use.
For example, the use of wollastonite is restricted because of
its relatively high cost, which is attributable to other major industrial uses for the material (ceramics, friction products, metallurgy, paint, and plastics) as well as the cost of transport
from isolated mined deposits to agricultural areas. Wollastonite is, however, a very effective source of fertilizer Si (Berthelsen et al., 2003; Korndorfer et al., 2005; Pereira and Cabral,
2005; Buck et al., 2011). Wollastonite is also a liming material
since its dissolution releases OH ions:
CaSiO3 + 3 H2O > Ca2+ + H4SiO4 + 2 OH
Diatomaceous earth is the fossilized remains of diatoms (phytoplankton that are enclosed in a cell wall made of silica) and
is found in sedimentary rock. While wollastonite solubility is
favored in the acidic pH range and therefore can be an effec-
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
www.plant-soil.com
838
Haynes
size and raising the pH during gel preparation (Vogelsberger

et al., 1992). Like diatomaceous earth and silica fume, its solubility is increased greatly above pH 9.0. It has been used
successfully in both paddy rice and wheat (Mecfel et al.,
2007; Gocke et al., 2013). Potassium silicate is manufactured
by calcining silica sand with K-carbonate and can be used as
a solid fertilizer product (Ma and Takahashi, 2002). Its high
solubility means it can be also used to supply plants with Si in
hydroponics or as foliar applications (Menzies et al., 1992;
Kanto et al., 2004).
As noted above, the total Si content of a fertilizer material
does not give any indication of its solubility or how much of
the Si is potentially plant-available. A number of different
chemical extractants for estimating available Si in silicate
fertilizer materials have been suggested (Kato and Owa,
1997a; 1997b; Haynes et al., 2013). These include 0.5 M HCl
(Savante et al., 1999), 0.5 M NH4-acetate (pH 4.8) (Hagihara,
1981), water/amberlite resin in H+ form (Kato and Owa,
1997b), and Na2CO3/NH4NO3 (Pereira et al., 2003). As already noted, dissolution of wollastonite and slag results in
consumption of H+ ions, and as pH rises further Si dissolution
is depressed. For this reason, acid extractants promote slag
dissolution (by removing OH from solution) and are believed
to extract unrealistically large amounts of Si (Imaizumi and
Yoshida, 1958; Kato and Owa, 1997a). The H+ resin method
used by Kato and Owa (1997b) buffers the extraction between pH 6 and 7 (similar values to those in soil). The
Na2CO3/NH4NO3 extractant was developed with the specific
aim of promoting dissolution of water-insoluble CaSiO3
(Pereira et al., 2003).
Working with a range of fertilizer Si materials and a range of
extractants, both Berthelsen et al. (2003) and Haynes et al.
(2013) concluded that there is no extractant that is reliable
over a range of Si sources. For example Na2CO3/NH4NO3 is
effective at extracting Si from wollastonite but ineffective for
slags, while the water/amberlite H+ resin method is effective
for slags but not for wollastonite (Haynes et al., 2013). Both
Pereira et al. (2003) and Berthelsen et al. (2003) demon-

strated that for a range of Si sources; the water/amberlite H+
resin method showed highest correlations with plant Si uptake. Nonetheless, in a similar experiment, Buck et al. (2011)
found the Na2CO3/NH4NO3 method was best correlated with
plant Si uptake. At present, most regulatory systems for Si
fertilizers use the total Si content of the materials.
Since wollastonite and slags are effective liming materials,
their application increases both soluble Si concentrations and
soil pH. This can result in a strong positive relationship between soluble or extractable Si and pH (Berthelsen et al.,
2003; Haynes et al., 2013). The relationship between pH and
CaCl2-extractable Si in a soil, when blast furnace slag was
applied at increasing rates to a Si-deficient soil, is shown in
Fig. 5. Such a relationship is often likely to be observed in
soils treated with varying amounts of wollastonite, blast furnace or steel slags. Adsorption of Si to soil colloids in the pH
range 56 is not strong, therefore any increased adsorption
caused by the increased pH is masked by the large increase
in Si supply.
Even where non-liming fertilizer Si sources (e.g., acidulated
wollastonite) are applied to soils, increasing pH can cause an
increase in Si solubility (Tavakkoli et al., 2011). In such cases,
decreased solubility at low pH may be a factor. For example,
silicic acid can readily combine with Al to form colloidal hydroxyaluminosilicates (HASs) (Farmer et al., 1979; Doucet et
al., 2001; Schneider et al., 2004). The precipitation of such
species has been forwarded as one mechanism by which addition of silicates to soils alleviates Al toxicity (Liang et al.,
2001; 2007). Doucet et al. (2001) showed that an excess of
Al over Si(OH)4 (as would typically occur in an acid soil) results in the formation of an imogolite-like phase termed HASA,
which has a Si:Al ratio of 0.5. When Si(OH)4 is present in excess (e.g., after Si application to a soil) then further reactions
occur to form HASB, which has a Si:Al ratio of 1.0. Both forms
are known to occur in soils (Doucet et al., 2001). The practical
significance of such reactions from the viewpoint of Si availability is that at soil pH values < 5.5 there are typically substantial concentrations of exchangeable and soluble Al
present, and added Si is likely to react with Al to form insoluble HASs. For example, addition of silicic acid to two acid
soils was shown by Matlou (2006) to lower exchangeable and
soil solution Al concentrations significantly.
Figure 5: Relationship between pH and CaCl2-extractable Si in a soil

treated with increasing rates of blast furnace slag. Redrawn from
Haynes et al. (2013).
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
www.plant-soil.com

not significantly influence tissue Si application while soil application of Ca-silicate was very effective.
Since it is the Si status of the plant that influences plant tolerance to biotic and abiotic stress, soil applications are generally more effective. Guevel et al. (2007), for instance, found
that soil applications of Si increased leaf Si concentrations
and reduced powdery mildew severity in wheat by as much
as 80% but foliar applications had a very small effect. Similar
observations were made by Liang et al. (2005) working with
powdery mildew on cucumber. In rice, brown spot incidence
was greatly reduced by soil application of Si, but there was
only a small effect of foliar-applied Si (Rezende et al., 2009).
It was suggested by Guevel et al. (2007) that foliar sprays of
K2SiO3 might inhibit powdery mildew severity by having osmotic effects on spores germinating on the leaf surface. Liang
et al. (2005) suggested that foliar Si applications do not elicit
physiological and biochemical/molecular mechanisms of host
resistance to pathogens that occur with soil applied applications. Thus, present evidence suggests that soil applications
of Si are required to achieve increased plant tolerance to biotic and abiotic stress and thus increased crop yields.
Irrigation water
The amount of Si supplied in irrigation water may well be an

important management consideration (Kingston, 2008). In
Hawaii, Fox et al. (1967) showed that the response of sugarcane to Si applications was markedly greater in areas irrigated with mountain runoff waters containing 2.5 mg Si L1
than where crops were irrigated with lower altitude groundwater containing 30 mg L1. Takahashi (2002) observed that
Si concentrations in irrigation waters in rice paddies in Japan
ranged between 1.9 and 28.9 mg L1. In average, 10.2 mg Si
L1 resulted in a supply of 142 kg Si ha1 (30% of the crop
needs) when irrigated with 4 ML ha1. Ma and Takahashi
(2002) also stressed the importance of Si input from irrigation
for paddy rice. Desplanques et al. (2006) measured an input
of 100 kg Si ha1 in irrigation water in a rice field in France.
They suggested that most of this Si originated from freshwater diatoms present in the irrigation water. This Si would
therefore be present as silica in the frustules of diatoms rather
than as soluble silicic acid, and the silica would need to dissolve before becoming plant-available. On the other hand, depletion of Si from irrigation water in some parts of Japan has
been attributed to rising densities of diatoms in reservoirs.
Flushes of diatom growth in reservoirs, caused by N and P
contamination of surface waters, results in removal of soluble
Si from the water and decreases in concentrations of soluble
Si in irrigation water downstream (Husnain et al., 2009).
Thus, the nature and magnitude of input of Si in irrigation
water needs further study. It should certainly be measured
and considered when considering applying Si fertilizers.
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-

erable primary and secondary minerals (since the return of
phytogenic Si will be limited). Globally, seven out of the 10
most important crops are Si-accumulator plants, so that in the
future crops other than rice and sugarcane (e.g., wheat, barley, soybeans, sugar beet, tomatoes) will very likely require Si
applications to maintain maximum yields. Since crops in the
field are constantly subjected to an array of stresses and plant
tolerance to many stress factors can be greatly improved by
Si applications, a lack of Si may well be contributing to a global yield decline which has been noted in recent times for a
range of crops. In addition, Si is now being promoted as an
environmentally-friendly plant, prophylactic for a very wide
range of crops, and soil applications have been shown to be
much more effective than foliar sprays. A much better understanding of the Si fertility of soils is therefore required.
There is a wide range of issues encompassing the solubility
and availability of Si in agricultural soils that need further research. Firstly, the relative importance of various solid phase
pools supplying the soil solution needs greater understanding. For example, the significance of adsorption/desorption
reactions is unclear since Si is preferentially adsorbed as
H3SiO4 (pK1 = 9.8) with an adsorption maximum at pH 9.8,
yet Si in soil solution, at common soil pH values of 56, is
present predominantly as H4SiO4. The fact that leaching of Si
from soils and its transport in streams to the sea are important
components of the biogeochemical cycling of Si suggests
that it is highly mobile in most soils. Limited research suggests that at soil pH values < 5.5, addition of Si can result in
the formation of insoluble hydroxyaluminosilicates (due to the
initially high soluble and exchangeable Al present), thus lowering Si availability. The significance of this reaction is, however, still unclear. The importance of phytogenic Si as a
source of soluble Si in agricultural soils also needs much
greater attention. A few studies have suggested that diminution of phytogenic Si is the main reason for the very low soluble Si concentrations in some agricultural soils. That is, phytogenic Si is removed at harvest rather than being returned to
the soil. Much more research is necessary to confirm the extent of this and to quantify the role of Si removal in harvested
crops as a factor in the accelerated desilication of agricultural
soils. The magnitude and form of inputs of Si in irrigation
water also needs greater consideration.
The effects of pH on the availability of soil Si needs greater
understanding due to the competing and interacting processes involved. Unlike litho-pedogenic minerals, in which Si
solubility is highest at low pH for phytogenic Si, solubility increases markedly with increasing pH. As already noted, likewise does specific adsorption of Si to hydrous oxide surfaces.
However, when fertilizer Si is applied as slag materials, soil
pH is also greatly increased since they are liming materials as
well as Si sources. As a result, both soluble Si and pH increase. Where non-liming Si sources are used, formation of
hydroxyaluminosilicates at low pH might limit Si availability.
Thus, the effect of pH on Si availability is, at present, unclear
and warrants further research.
www.plant-soil.com
840
Haynes
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.
References
Acquaye, D., Tinsley, J. (1965): Soluble Silica in Soils, in Hallsworth,
E. G., Crawford, D. V. (eds.): Experimental Pedology. Butterworth,
London, UK, pp. 126148.
Alexandre, A., Meunier, J.-D., Colin, F., Koud, J.-M. (1997): Plant
impact on the biogeochemical cycle of silicon and related weathering processes. Geochim. Cosmochim. Ac. 61, 677682.

Brunings, A. M., Datnoff, L. E., Ma, J. F., Mitani, N., Nagamura, Y.,
Rathinasabapathi, B., Kirst, M. (2009): Differential gene
expression of rice in response to silicon and rice blast fungas
Magnaporthe oryzae. Ann. Appl. Biol. 155, 161170.
Buck, G. B., Korndorfer, G. H., Datnoff, L. E. (2011): Extractors for
estimating plant available silicon from potential silicon fertilizer
sources. J. Plant Nutr. 34, 274282.
Carey, J. C., Fulweiler, R. W. (2012): The terrestrial silica pump.
PLoS ONE, DOI: 10.1371/journal.pone.0052932.
Cheong, W. Y., Ross, L., Cavalot, P. C. (1968): Silicon Status of Mauritius Soils. 1967 Annual report of the Mauritius Sugarcane Industry
Research Institute, Mauritius, pp. 7881.
Clarke, J. (2003): The occurrence and significance of biogenic opal in
the regolith. Earth Sci. Rev. 60, 175194.
Alexandre, A., Bouvet, M., Abbadie, L. (2011): The role of savannas

in the terrestrial Si cycle: a case study from Lamto, Ivory Coast.
Global Planet. Change 78, 162169.
Clements, H. F. (1965): Effects of silicate on the growth and leaf

freckle of sugar cane in Hawaii. Proc. Int. Soc. Sugar Cane
Technol. Soc. 12, 197215.
Ali, M. A., Lee, C. H., Kim, P. J. (2008): Effect of silicate fertilizer on

reducing methane emission during rice cultivation. Biol. Fertil.
Soils 44, 597604.
Clymans, W., Struyf, E., Govers, G., Vandevenne, F., Conley, D. J.

(2011): Anthropogenic impact on amorphous silica pools in
temperate soils. Biogeosciences 8, 22812293.
Ayres, R. S. (1966): Calcium silicate as a growth stimulant for

sugarcane on low silicon soils. Soil Sci. 101, 216227.
Cooke, J., Leishman, M. R. (2011): Is plant ecology more siliceous

than we realise? Trends Plant Sci. 16, 6168.
Barbosa Filho, M. P., Pfeilsticker Zimmermann, F. J., da Silva, O. F.

(2004): Influencia da escoria silicatada na acidez do solo e na
produtividade de graos do arroz de terras altas Cienc. Agrotec.
28, 323331 (in Portuguese).
Cornelis, J.-T., Ranger, J., Iserentant, A., Delvaux, B. (2010): Tree

species impact the terrestrial cycle of silicon through various
uptakes. Biogeochemistry 97, 231245.
Bartoli, F. (1983): The biogeochemical cycle of silicon in two

temperate forest ecosystems. Ecol. Bull. 35, 469476.
Berthelsen, S. (1997): Impact of Silicon on Sugarcane Productivity.
Yield Decline Joint Venture Working Group Meeting, 1012
September 1997, pp. 3640.
Berthelsen, S. (2000): An Assessment of the Silicon Status of Soils in
north Queensland and the Impact of Suboptimal Plant Available
Soil Silicon on Sugarcane Production Systems. MSc Thesis,
James Cook University, Townsville, Australia.
Cornelis, J.-T., Delvaux, B., Georg, R. B., Lucas, Y., Ranger, J.,
Opfergelt, S. (2011): Tracing the origin of dissolved silicon transferred from various soilplant systems towards rivers: a review.
Biogeosciences 8, 89112.
Correa, R. S. B., Moraes, J. C., Auad, M. Carvalho, G. A. (2005):
Silicon and acibenzolar-S-methyl as resistance inducers in
cucumber, against the whitefly Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae) biotype B. Neotrop. Entomol. 34, 429433.
Currie, H. A., Perry, C. C. (2007): Silica in plants: biological, biochemical and chemical studies. Ann. Bot. 100, 13831389.
Berthelsen, S., Noble, A. D., Garside, A. L. (2001): Silicon Research

Down Under: Past, Present and Future, in Datnoff, L. E., Snyder,
G. H., Korndorfer, G. H. (eds.): Silicon in Agriculture. Elsevier,
Amsterdam, The Netherlands, pp. 241255.
Desplanques, V., Cary, L., Mouret, J.-C., Trolard, F., Bourrie, G.,
Grauby, O., Meunier, J.-D. (2006): Silicon transfers in a rice field in
Camargue (France). J. Geochem. Explor. 88, 190193.
Berthelsen, S., Noble, A. D., Kingston, G., Hurney, A. P., Rudd, A.,
Garside, A. L. (2003): Improving Yield and ccs in Sugarcane
Through the Application of Silicon Based Amendments. Final
Report, Sugar Research and Development Corporation, Project
CLW009, May 2003.
Dobermann, A., Fairhurst, T. H. (2000): RiceNutrient Disorders and

Nutrient Management. Potash and Phosphate Institute, Singapore.
Berthelsen, S., Korndorfer, G. (2005): Methods for Si analysis in

plant, soil and fertilizers. Proceedings of the 3rd International
Conference Silicon in Agriculture, 2226 October 2005,
Uberlkander, Brazil, pp. 8289.
Bharwana, S. A., Ali, S., Farooq, M. A., Iqbal, N., Abbas, F., Ahmad,
M. S. A. (2013): Alleviation of lead toxicity by silicon is related to
elevated photosynthesis, antioxidant enzymes suppressed lead
uptake and oxidative stress in cotton. J. Bioremed. Biodeg. 4, 187.
Blecker, S. W., McCulley, R. L., Chadwick, O. A., Kelly, E. F. (2006):
Biologic cycling of silica across a grassland bioclimosequence.
Global Biogeochem. Cy. 20, DOI: 10.1029/2006GB002690.
Bowden, J. W., Posner, A. M., Quirk, J. P. (1980): Adsorption and
Charging Phenomena in Variable Charge Soils, in Theng, B. K. G.
(ed.): Soils of Variable Charge. New Zealand Society of Soil
Science, Lower Hutt, New Zealand, pp. 147166.
Bowen, P. A., Menzies, J. G., Ehret, D. L. (1992): Soluble silicon
sprays inhibit powdery mildew development on grape leaves. J.
Am. Soc. Hortic. Sci. 117, 906912.
Dietzel, M. (2000): Dissolution of silicates and the stability of polysilicic acid. Geochim. Cosmochim. Ac. 64, 32753281.
Doucet, F. J., Schneider, C., Bones, S. J., Kretchmer, A., Moss, I.,
Tekely, P., Exley, C. (2001): The formation of hydroxyaluminosilicates of geochemical and biological significance. Geochim.
Cosmochim. Acta 65, 24612467.
Dove, P. M. (1995): Kinetic and thermodynamic controls on silica
reactivity in weathering environments. Rev. Mineral. Geochem. 31,
235290.
Dove, P. M., Han, N., Wallace, A. F., De Yoreo, J. J. (2008): Kinetics
of amorphous silica dissolution and the paradox of the silica polymorphs. PNAS 22, 99039908.
Drees, L. R., Wilding, L. P., Smeck, N. E., Senkayi, A. L. (1989): Silica
in Soils: Quartz and Disordered Silica Polymorphs, in Dixon, J. B.,
Weed, S. B. (eds.): Minerals in the Soil Environments. Soil Science
Society of America, Madison, WI, USA, pp. 914974.
Dufey, I., Gheysens, S., Ingabire, A., Lutts, S., Bertin, P. (2013):
Silicon application in cultivated rices (Oryza sativa L. and Oryza
glaberrima Steud) alleviates iron toxicity symptoms through the
reduction in iron concentration in the leaf tissue. J. Agron. Crop
Sci. 200, 132142.
www.plant-soil.com
Elawad, S. H., Gascho, G. J., Street, J. J. (1982): Response of

sugarcane to silicate source and rate. I. Growth and yield. Agron.
J. 74, 481484.
Gomes, F. B., de Moraes, J. C., dos Santos, C. D., Goussain, M. M.

(2005): Resistance induction in wheat plants by silicon and aphids.
Sci. Agr. 62, 547551.
Epstein, E. (1999): Silicon. Annu. Rev. Plant Phys. 50, 641664.
Guevel, M.-H., Menzies, J. G., Belanger, R. R. (2007): Effect of root

and foliar applications of soluble silicon on powdery mildew control
and growth of wheat plants. Eur. J. Plant Pathol. 119, 429436.
Epstein, E. (2001): Silicon in Plants: Facts vs. Concepts, in Datnoff,

L. E., Snyder, G. H., Korndorfer, G. H. (eds.): Silicon in Agriculture.
Studies in Plant Science, Vol. 8, Elsevier, Amsterdam, The Netherlands, pp. 115.
Farmer, V. C., Fraser, A. R., Tait, J. M. (1979): Characterization of the
chemical structures of natural and synthetic aluminosilicate gels
and sols by infrared spectroscopy. Geochim. Cosmochim. Ac. 43,
14171420.
Farmer, V. C., Delbos, E., Miller, J. D. (2005): The role of phytolith
formation and dissolution in controlling concentrations of silica in
soil solutions and streams. Geoderma 127, 7179.
Fauteux, F., Chain, F., Belzile, F., Menzies, J. G., Belanger, R. R.
(2006): The protective role of silicon in the Arabidopsis-powdery
mildew pathosystem. P. Natl. Acad. Sci. USA 103, 1755417559.
Fox, R. L., Silva, J. A., Younge, O. R., Plucknett, D. L., Sherman, G.
D. (1967): Soil and plant silicon response by sugarcane. Soil Sci.
Soc. Amer. J. 31, 775779.
Foy, C. D. (1992): Soil chemical factors limiting plant growth. Adv.
Soil Sci. 19, 97149.
Fraysse, F., Pokrovsky, O. S., Schott, J., Meunier, J.-D. (2006):
Surface properties, solubility and dissolution kinetics of bamboo
phytoliths. Geochim. Cosmochim. Ac. 70, 19301951.
Fraysse, F., Pokrovsky, O. S., Schott, J., Meunier, J.-D. (2009):
Surface chemistry and reactivity of plant phytoliths in aqueous
solutions. Chem. Geol. 258, 197206.
Fraysse, F., Pokrovsky, O. S., Meunier, J.-D. (2010): Experimental
study of terrestrial plant litter interaction with aqueous solutions.
Geochim. Cosmochim. Ac. 74, 7084.
Frings, P. J., Clymans, W., Jeppesen, E., Lauridsen, T. L., Struyf, E.,
Conley, D. J. (2014): Lack of steady-state in the global biogeochemical Si cycle: emerging evidence from lake Si sequestration.
Biogeochemistry 117, 255277.
Gallez, A., Herbillon, A. J., Juo, A. S. R. (1977): Characteristics of
silica sorption and solubility as parameters to evaluate the surface
properties of tropical soils: I. The index of silica reactivity. Soil Sci.
Soc. Am. J. 41, 11461150.
Gascho, G. J. (2001): Silicon Sources for Agriculture, in Datnoff, L.
E., Snyder, G. H., Korndorfer, G. H. (eds.): Silicon in Agriculture.
Studies in Plant Science, Vol. 8, Elsevier, Amsterdam, The Netherlands, pp. 197207.
Gascho, G. J., Andreis, H. J. (1974): Sugarcane response to calcium
silicate slag applied to organic and sand soils. Proc. Int. Soc.
Sugar Cane Technol. 15, 543551.
Gerard, F., Mayer, K. U., Hodson, M. J., Ranger, J. (2008): Modelling
the biogeochemical cycle of silicon in soils: application to a
temperate forest ecosystem. Geochim. Cosmochim. Ac. 72, 741
758.
Ghareeb, H., Bozso, Z., Ott, P. G., Repenning, C., Stahl, F., Wydra,
K. (2011): Transcriptome of silicon-induced resistance against
Ralstonia solanacearum in the silicon non-accumulator tomato
implicates priming effect. Physiol. Mol. Plant P. 75, 8389.
Gocke, M., Liang, W., Sommer, M., Kuzyakov, Y. (2013): Silicon
uptake by wheat: effects of Si pools and pH. J. Plant Nutr. Soil Sci.
176, 551560.
Goldberg, S., Glaubig, R. A. (1988): Boron and silicon adsorption on
an aluminium oxide. Soil Sci. Soc. Am. J. 52, 8791.
Guntzer, F., Keller, C., Meunier, J.-D. (2012a): Benefits of plant silicon
for crops: a review. Agron. Sustain. Dev. 32, 201213.
Guntzer, F., Keller, C., Poulton, P. R., McGrath, S. P., Meunier, J.-D.
(2012b): Long-term removal of wheat straw decreases soil amorphous silica at Broadbalk, Rothamsted. Plant Soil 351, 173184.
Gu, H.-H., Zhan, S.-S., Wang, S.-Z., Tang, Y.-T., Chaney, R. L., Fang,
X.-H., Cai, X.-D., Qiu, R.-L. (2012): Silicon-mediated amelioration
of zinc toxicity in rice (Oryza sativa L.) seedlings. Plant Soil 350,
193204.
Hagihara, H. H. (1981): Laboratory Evaluation of Residual Properties
of Siliceous Materials. Agronomy Report 40, Hawaiian Sugar
Planters Association. Honolulu, HI, USA.
Haynes, R. J. (1984): Lime and phosphate in the soil-plant system.
Adv. Agron. 37, 249315.
Haynes, R. J., Belyaeva, O. N., Kingston, G. (2013): Evaluation of
industrial wastes as sources of fertilizer silicon using chemical
extractions and plant uptake. J. Plant Nutr. Soil Sci. 176, 238248.
Haysom, M. B. C., Chapman, L. S. (1975): Some aspects of the
calcium silicate trials at Mackay. Proc. Aust. Sugar Cane Technol.
42, 117122.
Heinai, H., Saigusa, M. (2006): Silicon availability of nursery bed
soils and effects of silicon fertilizer applied on the growth and
silicon uptake of rice (Oryza sativa L.) seedlings. Soil Sci. Plant
Nutr. 77, 4146.
Herbillon, A. J., Gallez, A., Juo, A. S. R. (1977): Characteristics of
silica sorption and solubility as parameters to evaluate the surface
properties of tropical soils: II. The index of silica saturation. Soil
Sci. Soc. Am. J. 41, 11511154.
Hingston, F. J., Raupach, M. (1967): The reaction between monosilicic acid and aluminium hydroxide. I. Kinetics of adsorption of
silicic acid by aluminium hydroxide. Aust. J. Soil Res. 5, 295309.
Hingston, F. J., Posner, A. M., Quirk, J. P. (1972): Anion adsorption
by goethite and gibbsite. I. The role of the proton in determining
adsorption envelopes. Eur. J. Soil Sci. 23, 177192.
Hohn, A., Sommer, M., Kaczorek, D., Schalitz, G., Breuer, J. (2008):
Silicon fractions in Histosols and Gleysols of a temperate
grassland site. J. Plant Nutr. Soil Sci. 171, 409418.
Hurney, A. P. (1973): A progress report on calcium silicate investigations. Proc. Qld. Soc. Sugar Cane Technol. 40, 109113.
Husnain, H., Wakatsuki, T., Masunaga, T. (2009): Dissolved silica
dynamics and phytoplankton population in Citarum watershed,
Indonesia. J. Food Agric. Environ. 7, 655661.
Imaizumi, K., Yoshida, S. (1958): Edaphological studies on silicon
supplying power of paddy soils. Bull. Natl. Inst. Agric. Sci. (Jpn.)
B8, 261304.
Juo, A. S. R., Sanchez, P. A. (1986): Soil nutritional aspects with a
view to characterize upland rice environments, in IRRI (ed.):
Progress In Upland Rice Research, International Rice Research
Institute, Los Banos, Philippines, pp. 8591.
Kanamugire, A. (2007): A comparison of soil extraction methods for
predicting the silicon requirements for sugarcane. MSc thesis,
University of KwaZulu-Natal, South Africa.
Kanamugire, A., Meyer, J. H., Haynes, R. J., Naidoo, G., Keeping,
M. G. (2006): An assessment of soil extraction methods for
predicting the silicon requirement of sugarcane. Proc. South Afr.
Sugar Technol. Assoc. 80, 287290.
www.plant-soil.com
842
Haynes
Kang, Y. S., Jung, Y.-T. (2002): Research on agricultural utilization of

silicon in Korea. Proceedings of the 2nd International Conference
Silicon in Agriculture, 2227 August 2002, Tsuruoka, Japan, pp.
2226.
Kanto, T., Miyoshi, A., Ogawa, T., Maekawa, K., Aino, M. (2004):
Suppressive effect of potassium silicate on powdery mildew of
strawberry in hydroponics. J. Gen. Plant Pathol. 70, 207211.
Karathanasis, A. D. (2002): Mineral Equilibria in Environmental Soil
Systems, in Dixon, J. B., Schulze, D. G. (eds.): Soil Mineralogy
with Environmental Applications. Soil Science Society of America,
Madison, WI, USA, pp. 109151.
Karathanasis, A. D. (2006): Clay Minerals: Weathering and Alteration
of, in Lal, R. (ed.): Encyclopedia of Soil Science, Vol. 2. CRC
Press, Boca Raton, FL, USA, pp. 281286.
Kato, N., Owa, N. (1997a): Dissolution of slag fertilizers in a paddy
soil and Si uptake by rice plant. Soil Sci. Plant Nutr. 43, 329341.
Kato, N., Owa, N. (1997b): Evaluation of Si availability in slag fertilizers by an extraction method using a cation exchange resin. Soil
Sci. Plant Nutr. 43, 351359.
Keeping, M. G., Kvedaras, O. L. (2008): Silicon as a plant defence
against insect herbivory: response to Massey, Ennos and Hartley.
J. Anim. Ecol. 77, 631633.

Liang, Y., Sun, W., Zhu, Y.-G., Christie, P. (2007): Mechanisms of
silicon-mediated alleviation of abiotic stresses in higher plants: a
review. Environ. Poll. 147, 422428.
Lucas, Y., Luizao, F. J., Chauvel, A., Roullier, J., Nahon, D. (1993):
The relation between biological activity of a rainforest and mineral
composition of the soils. Science 260, 521523.
Ma, J. F., Takahashi, E. (2002): Soil, Fertilizer and Plant Silicon
Research in Japan. Elsevier, Amsterdam, The Netherlands.
Ma, J. F., Yamaji, N. (2006): Silicon uptake and accumulation in higher plants. Trends Plant Sci. 11, 392397.
Maksimovic, J. D., Mojovic, M., Maksimovic, V., Romheld, V., Nikolic,
M. (2012): Silicon ameliorates manganese toxicity in cucumber by
decreasing hydroxyl radical accumulation in the leaf apoplast. J.
Exp. Bot. 63, 24112420.
Matichenkov, V. V., Bocharnikova, E. A. (2001): The Relationship between Silicon and Soil Physical and Chemical Properties, in
Datnoff, L. E., Snyder, G. H., Korndorfer, G. H. (eds.): Silicon in
Agriculture. Studies in Plant Science, Vol. 8, Elsevier, Amsterdam,
The Netherlands, pp. 209219.
Matlou, M. G. (2006): A comparison of soil and foliar applied silicon
on nutrient availability and plant growth and soil-applied silicon on
phosphorus availability. MSc thesis, University of KwaZulu-Natal,
South Africa.
Keller, C., Guntzer, F., Barboni, D., Labreuche, J., Meunier, J.-D.
(2012): Impact of agriculture on the Si biogeochemical cycle: input
from phytolith studies. C. R. Geoscience 344, 739746.
McKeague, J. A., Cline, M. G. (1963): Silica in soils. Adv. Agron. 15,

339396.
Kingston, G. (2008): Silicon fertilizersRequirement and field experiences. Proceedings of the 4th International Conference Silicon in
Agriculture, 2631 October 2008, Wildcoast, South Africa, pp. 82
89.
Mecfel, J., Hinke, S., Goedel, W. A., Marx, G., Fehlhaber, R.,
Baucker, E., Wienhaus, O. (2007): Effect of silicon fertilizers on
silicon accumulation in wheat. J. Plant Nutr. Soil Sci. 170, 769
772.
Kingston, G., Berthelsen, S., Hurney, A. P., Rudd, A., Noble, A. D.

(2005): Impact of calcium silicate amendments on sugarcane yield
and soil properties in Queensland, Australia. Proceedings of the
3rd International Conference Silicon in Agriculture, 2226 October
2005, Uberlkander, Brazil, pp. 107117.
Meena, V. D., Dotaniya, M. L., Coumar, V., Rajendiran, S., Ajay,

Kundu, S., Rao, A. S. (2014): A case for silicon fertilization to
improve crop yields in tropical soils. P. Natl. Acad. Sci., India,
Sect. B Biol. Sci. DOI 10.1007/s40011-013-0270-y.
Korndorfer, G. H., Lepsch, I. (2001): Effect of Silicon on Plant Growth

and Crop Yield, in Datnoff, L. E., Snyder, G. H., Korndorfer, G. H.
(eds.): Silicon in Agriculture. Studies in Plant Science, Vol. 8,
Elsevier, Amsterdam, The Netherlands, pp. 133147.
Korndorfer, G. H., Nolla, A., Ramos, L. A. (2005): Available silicon in
tropical soils and crop yield. Proceedings of the 3rd International
Conference Silicon in Agriculture, 2226 October 2005,
Uberlkander, Brazil, pp. 7281.
Korndorfer, G. H., Pereira, H. S. (2011): Silicon testing, silicon fertilizer manufacturing techniques and standards. Proceedings of the
5th International Conference Silicon in Agriculture, 2226 October
2005, 1318 September 2011, Beijing, China, pp. 8998.
Kvedaras, O. L., Keeping, M. G. (2007): Silicon impedes stalk penetration by the borer Eldana saccharina in sugarcane. Entomol.
Exp. Appl. 125, 103110.
Kvedaras, O. L., An, M., Choi, Y. S., Gurr, G. M. (2010): Silicon
enhances natural enemy attraction and biological control through
induced plant defences. Bull. Entomol. Res. 100, 367371.
Menzies, J., Bowen, P., Ehret, D., Glass, A. D. M. (1992): Foliar applications of potassium silicate reduce severity of powdery mildew on
cucumber, muskmelon and zucchini squash. J. Am. Soc. Hortic.
Sci. 117, 902905.
Meunier, J.-D., Kirman, S., Strasberg, D., Nicolini, E., Delcher, E.,
Keller, C. (2010): The output and bio-cycling of Si in a tropical rain
forest developed on young basalt flows (La Reunion Island).
Geoderma 159, 431439.
Meyer, J. H., Keeping, M. G. (2001): Past, Present and Future
Research of the Role of Silicon for Sugarcane in Southern Africa,
in Datnoff, L. E., Snyder, G. H., Korndorfer, G. H. (eds.): Silicon in
Agriculture. Studies in Plant Science, Vol. 8, Elsevier, Amsterdam,
The Netherlands, pp. 257275.
Miles, N., Manson, A. D., Rhodes, R., van Antwerpen, R., Weigel, A.
(2014): Extractable silicon in soils of the South African sugar
industry and relationships with crop uptake. Commun. Soil Sci.
Plant Anal. (in press)
Mitani, N., Ma, J. F. (2005): Uptake system of silicon in different plant
species. J. Exp. Bot. 56, 12551261.
Laing, M. D., Gatarayiha, M. C., Adandonon, A. (2006): Silicon use

for pest control in agriculture: a review. Proc. South Afr. Sugar
Technol. Assoc. 80, 278286.
Mizuochi, T. (2002): Role of silicon in potassium silicate fertilizer.

Proceedings of the 2nd International Conference Silicon in Agriculture, 2227 August 2002, Tsuruoka, Japan, pp. 191194.
Liang, Y., Yang, C., Shi, H. (2001): Effects of silicon on growth and
mineral composition of barley grown under toxic levels of
aluminum. J. Plant Nutr. 24, 229243.
Nguyen, M. N., Dultz, S., Guggenberger, G. (2014): Effects of

pretreatment and solution chemistry on solubility of rice-straw
phytoliths. J. Plant Nutr. Soil Sci. 177, 349359.
Liang, Y. C., Sun, W. C., Si, J., Romheld, V. (2005): Effects of foliarand root-applied silicon on the enhancement of induced resistance
to powdery mildew in Cucumis sativus. Plant Pathol. 54, 678685.
Nhan, P. P., Hai, N. T. (2013): Amelioration of aluminium toxicity on

OM4900 rice seedlings by sodium silicate. Afric. J. Plant Sci. 7,
208212.
www.plant-soil.com
Owa, N. (2002): Silicate fertilizers in Japan. Proceedings of the 2nd

International Conference Silicon in Agriculture, 2227 August
2002, Tsuruoka, Japan, pp. 175180.
Sommer, M., Kaczorek, D., Kuzyakov, Y., Breuer, J. (2006): Silicon

pools and fluxes in soils and landscapesa review. J. Plant Nutr.
Soil Sci. 169, 310329.
Oya, K., Kina, K. (1989): Fertility studies on soils of Okinawa in

relation to sugarcane production. 3. pH and soluble silica in dark
red soils. Sci. Bull. Coll. Agric. Univ. Ryukyus, Okinawa 36, 2531.
Sommer, M., Jochheim, H., Hohn, A., Breuer, J., Zagorski, Z., Busse,
J., Barkusky, D., Meier, K., Puppe, D., Wanner, M., Kaczorek, D.
(2013): Si cycling in a forest biogeosystemthe importance of
transient state of biogenic Si pools. Biogeosciences 10, 4991
5007.
Oya, K., Kuroshio, K., Hokama, Y. (1989): Fertility studies on soils of

Okinawa in relation to sugarcane production. 2. Soil pH and
soluble silica. Sci. Bull. Coll. Agric. Univ. Ryukyus, Okinawa 36,
1923.
Parr, J. F., Sullivan, L. A. (2005): Soil carbon sequestration in phytoliths. Soil Biol. Biochem. 37, 117124.
Pereira, H. S., Cabral, N. B. (2005): Slag as a silicon source: application criterion. Proceedings of the 3rd International Conference
Silicon in Agriculture, 2226 October 2005, Uberlkander, Brazil,
pp. 90101.
Street-Perrott, F. A., Barker, P. A. (2008): Biogenic silica: a neglected

component of the coupled global continental biogeochemical
cycles of carbon and silica. Earth Surf. Process. Landforms 33,
14361457.
Struyf, E., Smis, A., Van Damme, S., Meire, P., Conley, D. J. (2009):
The global biogeochemical silicon cycle. Silicon 1, 207213.
Pereira, H. S., Korndorfer, G. H., Moura, W. F., Correa, G. F. (2003):

Extratores de silcio disponvel em escorias e fertilizantes. Rev.
Bras. Cienc. Solo 27, 265274 (in Portuguese).
Struyf, E., Smis, A., Van Damme, S., Garnier, J., Govers, G., Van
Wesemael, B., Conley, D. J., Batelaan, O., Frot, E., Clymans, W.,
Vandevenne, F., Lancelot, C., Goos, P., Meire, P. (2010): Historical
land use change has lowered terrestrial silica mobilization. Nat.
Comm. 1, 17.
Raid, R. N., Anderson, D. L., Ulloa, M. F. (1992): Influence of cultivar

and soil amendment with calcium silicate slag on foliar disease development and yield of sugarcane. Crop Prot. 11, 8488.
Sumida, H. (2002): Plant Available Silicon in Paddy Soils.

Rezende, D. C., Rodrigues, F. A., Carre-Missio, V., Schurt, D. A.,

Kawamura, I. K., Korndorfer, G. H. (2009): Effect of root and foliar
applications of silicon on brown spot development in rice.
Australas. Plant Path. 38, 6773.
Takahashi, E. (2002): An Introduction to Silicon Research in Japan.

Rovner, I. (1971): Potential of opal phytoliths for use in paleoecological reonstruction. Quatern. Res. 1, 343359.
Sacala, E. (2009): Role of silicon in plant resistance to water stress.
J. Elementol. 14, 619630.
Saccone, L., Conley, D. J., Koning, E., Sauer, D., Sommer, M.,
Kaczorek, D., Blecker, S. W., Kelly, E. F. (2007): Assessing the
extraction and quantification of amorphous silica in soils of forest
and grassland ecosystems. Eur. J. Soil Sci. 58, 14461459.
Saccone, L., Conley, D. J., Likens, G. E., Bailey, S. W., Buso, D. C.,
Johnson, C. E. (2008): Factors that control the range and variability
of amorphous silica in soils in the Hubbard Brook Experimental
Forest. Soil Sci. Soc. Am. J. 72, 16371644.
Santos, G. M., Alexandre, A., Coe, H. H. G., Reyerson, P. E.,
Southon, J. R., De Carvalho, C. N. (2010): The phytolith 14C
puzzle: a tale of background determinations and accuracy tests.
Radiocarbon 52, 113128.
Sauer, D., Saccone, L., Conley, D. J., Herrmann, L., Sommer, M.
(2006): Review of methodologies for extracting plant-available and
amorphous Si from soils and aquatic sediments. Biogeochemistry
80, 89108.
Savante, N. K., Snyder, G. H., Datnoff, L. E. (1997): Silicon
management and sustainable rice production. Adv. Agron. 58,
151199.
Savante, N. K., Korndorfer, G. H., Datnoff, L. E., Snyder, G. H.
(1999): Silicon nutrition and sugarcane production: a review. J.
Plant Nutr. 22, 18531903.
Schneider, C., Doucet, F., Strekopytov, S., Exley, C. (2004): The solubility of an hydroxyaluminosilicate. Polyhedron 23, 31853191.
Seyfferth, A. L., Kocar, B. D., Lee, J. A., Fendorf, S. (2013): Seasonal
dynamics of dissolved silicon in a rice cropping system after straw
incorporation. Geochim. Cosmochim. Ac. 123, 120133.
Smithson, F. (1956): Plant opal in soil. Nature 178, 107108.
Snyder, G. H. (1991): Development of a silicon soil test for Histosolgrown rice. EREC Res. Rep., EV-1991-2. Univ. Florida, Belle
Glade, FL, USA.
Tavakkoli, E., Lyons, G., English, P., Guppy, C. N. (2011): Silicon

nutrition of rice is affected by soil pH, weathering and silicon fertilisation. J. Plant Nutr. Soil Sci. 174, 437446.
Tripathi, D. K., Singh, V. P., Gangwar, S., Prasad, S. M., Maurya, J.
N., Chauhan, D. K. (2014): Role of Silicon in Enrichment of Plant
Nutrients and Protection from Biotic and Abiotic Stresses, in
Ahmad, P. (ed.): Improvement of Crops in the Era of Climatic
Changes, Vol. 1. Springer, New York, NY, USA, pp. 3956.
Van Bennekom, A. J., Buma, A. G. J., Nolting, R. F. (1991): Dissolved
aluminium in the Weddell-Scotia Confluence and effect of Al on the
dissolution of biogenic silica. Mar. Chem. 35, 423434.
Van Bockhaven, J., De Vleesschauwer, D., Hofte, M. (2013):
Towards establishing broad-spectrum disease resistance in plants:
silicon leads the way. J. Exp. Bot. 64, 12811293.
Vandevenne, F., Struyf, E., Clymans, W., Meire, P. (2012): Agricultural
silica harvest: have humans created a new loop in the global silica
cycle? Front. Ecol. Environ. 10, 243248.
Vogelsberger, W., Seidel, A., Rudakoff, G. (1992): Solubility of silica
gel in water. J. Chem. Soc. Faraday Trans. 88, 473476.
Voogt, W., Sonnenveld, C. (2001): Silicon in Horticultural Crops
Grown in Soil-Less Culture, in Datnoff, L. E., Snyder, G. H., Korndorfer, G. H. (eds.): Silicon in Agriculture. Studies in Plant Science,
Vol. 8, Elsevier, Amsterdam, The Netherlands, pp. 115131.
Wang, H., Li, C., Liang, Y. (2001): Agricultural Utilization of Silicon in
China, in Datnoff, L. E., Snyder, G. H., Korndorfer, G. H. (eds.):
Silicon in Agriculture. Studies in Plant Science, Vol. 8, Elsevier,
Amsterdam, The Netherlands, pp. 343358.
Wang, J. J., Dodla, S. K., Henderson, R. E. (2004): Soil silicon
extractability with seven selected extractants in relation to colorimetric and ICP determination. Soil Sci. 169, 861870.
Wang, M. (2005): Effect of complete silicon fertilizer on rice yield.
Acta Agric. Shanghai 21, 7173.
Wedepohl, K. H. (1995): The composition of the continental crust.
Geochim. Cosmochim. Ac. 59, 12171232.
White, A. F. (1995): Chemical weathering rates of silicate minerals in
soils. Rev. Mineral Geochem. 31, 407461.
www.plant-soil.com
844
Haynes
White, A. F., Vivit, D. V., Shulz, M. S., Bullen, T. D., Evett, R. R.,
Aagarwal, J. (2012): Biogenic and pedogenic controls on Si distributions and cycling in grasslands of the Santa Cruz soil chronosequence, California. Geochim. Cosmochim. Ac. 94, 7294.

Ye, M., Song, Y., Long, J., Wang, R., Baerson, S. R., Pan, Z.,
Zhu-Salzman, K., Xie, J., Cai, K., Luo, S., Zeng, R. (2013): Priming
of jasmonate-mediated antiherbivore defense responses in rice
by silicon. P. Natl. Acad. Sci. USA 110, DOI: 10.1073/
pnas.1305848110. Epub 2013 Sep 3.
Wickramasinghe, D. B., Rowell, D. L. (2006): The release of silicon

from amorphous silica and rice straw in Sri Lankan soils. Biol.
Fertil. Soils 42, 231240.
Yoshida, S. (1981): Fundamentals of Rice Crop Science. International Rice research Institute, Philippines.
Wilding, L. P., Hallmark, C. T., Smeck, N. E. (1979): Dissolution and

stability of biogenic opal. Soil Sci. Soc. Am. J. 43, 800802.
Zhu, Y., Gong, H. (2014): Beneficial effects of silicon on salt and

drought tolerance in plants. Agron. Sustain. Dev. 34, 455472.
www.plant-soil.com

A Contemporary Overview of Silicon Availability in Agricultural Soils

Загружено:

Сведения о документе

Авторское право

Доступные форматы

Поделиться этим документом

Поделиться или встроить документ

Параметры публикации

Этот документ был вам полезен?

Это неприемлемый материал?

Авторское право:

Доступные форматы

A Contemporary Overview of Silicon Availability in Agricultural Soils

Загружено:

Авторское право:

Доступные форматы

J. Plant Nutr. Soil Sci.

2014, 177, 831844

A contemporary overview of silicon availability in agricultural soils

Accepted September 19, 2014

Silicon (Si) is the second most abundant element of the

agricultural systems is creating a new removal mechanism for

* Correspondence: Prof. R. J. Haynes; e-mail: r.haynes1@uq.edu.au

2014 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

J. Plant Nutr. Soil Sci. 2014, 177, 831844

Silicon status of soils

A major consequence of the chemical weathering of primary

of their production is on highly weathered Ultisols and Oxisols

Silicate adsorbed / mol cm2

Figure 1: Relationship between pH and CaCl2-extractable Si for 112

2014 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

Figure 2: Effect of pH and ionic strength on adsorption of silicate by

J. Plant Nutr. Soil Sci. 2014, 177, 831844

Silicon uptake and use by crops

2014 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

Silicon availability in agricultural soils 833

As already noted, Si accumulates in plants as silica deposits

crease with depth (Alexandre et al., 1997; Saccone et al.,

2014 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

J. Plant Nutr. Soil Sci. 2014, 177, 831844

J. Plant Nutr. Soil Sci. 2014, 177, 831844

As discussed above, phytogenic Si is now recognized as one

Silicon availability in agricultural soils 835

Log R / mol Si g1 day1

R / mol Si cm2 sec1

2014 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

J. Plant Nutr. Soil Sci. 2014, 177, 831844

crease the solubilization and plant-availability of phytogenic

dispersion of soil particles (Berthelsen and Korndorfer, 2005)

Ammonium- and Na-acetates (buffered at low pH with acetic

There are three key parameters that are likely to be important

2014 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

Citric acid has been suggested by some workers (Acquaye

J. Plant Nutr. Soil Sci. 2014, 177, 831844

Silicon availability in agricultural soils 837

Extraction ratio and period

Suggested critical level Reference

Fox et al. (1967);

Haysom and Chapman

0.5 M NH4-acetate (pH 4.8)

Fox et al. (1967)

0.5 M acetic acid

0.18 Na-acetate (plus 0.87 M acetic acid, pH 4.0)

Imaizumi and Yoshida (1958)

0.005 M sulfuric acid

0.025 M sulfuric acid

1:10 for 20 min

Meyer and Keeping (2001);

0.04 M Na2HPO4 (pH 6.2)

Heinai and Saigusa (2006)

There is a range of silicate-containing materials suitable for

2014 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

size and raising the pH during gel preparation (Vogelsberger

J. Plant Nutr. Soil Sci. 2014, 177, 831844

Figure 5: Relationship between pH and CaCl2-extractable Si in a soil

2014 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

J. Plant Nutr. Soil Sci. 2014, 177, 831844

The amount of Si supplied in irrigation water may well be an

2014 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

Silicon availability in agricultural soils 839