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Silicons Role in Abiotic

and Biotic Plant Stresses
Daniel Debona,1 Fabrcio A. Rodrigues,1
and Lawrence E. Datnoff 2
Department of Plant Pathology, Laboratory of Host-Pathogen Interaction, Vicosa Federal
University, Vicosa, Minas Gerais State, Brazil, 36570-900; email:
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Department of Plant Pathology & Crop Physiology, Louisiana State University Agricultural
Center, Baton Rouge, Louisiana, 70803; email:

Annu. Rev. Phytopathol. 2017. 55:4.14.23 Keywords

The Annual Review of Phytopathology is online at disease management, drought, foliar diseases, host defense mechanisms,
metal toxicity, pest control, plant nutrition, salt stress, soilborne diseases
035312 Abstract
Copyright  c 2017 by Annual Reviews. Silicon (Si) plays a pivotal role in the nutritional status of a wide variety
All rights reserved
of monocot and dicot plant species and helps them, whether direct or in-
directly, counteract abiotic and/or biotic stresses. In general, plants with a
high root or shoot Si concentration are less prone to suffer from pest at-
tack and exhibit enhanced tolerance to abiotic stresses such as drought, low
temperature, or metal toxicity. However, the most notable effect of Si is the
reduction in the intensities of a number of seedborne, soilborne, and foliar
diseases in many economically important crops that are caused by biotrophic,
hemibiotrophic, and necrotrophic plant pathogens. The reduction in disease
symptom expression is due to the effect of Si on some components of host
resistance, including incubation period, lesion size, and lesion number. The
mechanical barrier formed by the polymerization of Si beneath the cuticle
and in the cell walls was the first proposed hypothesis to explain how this
element reduced the severity of plant diseases. However, new insights have
revealed that many plant species supplied with Si have the phenylpropanoid
and terpenoid pathways potentiated and have a faster and stronger tran-
scription of host defense genes and higher activities of defense enzymes.
Photosynthesis and the antioxidant metabolism involved in the removal of
reactive oxygen species are improved for Si-supplied plants. Although the
current understanding of how this overlooked element affects plants against
pathogen infections, pest attacks, and abiotic stresses has advanced, the exact
mechanism(s) by which it modulates plant physiology through the poten-
tiation of host defense mechanisms still needs further investigation at the
genomic, metabolomic, and proteomic levels.


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Silicon (Si), a tetravalent metalloid, is the second most abundant element in the Earths crust
after oxygen (43). Most soils contain Si concentrations ranging from 14 to 20 mg Si/L (10).
Concentrations of this magnitude are common for other inorganic nutrients, such as sulfur (S),
potassium (K), and calcium (Ca), and are in excess of phosphorus (P) concentrations in the soil
solution (10, 43, 160). Dissolution of Si from soil minerals is a slow process and its adsorption
or resorption by the soil particles along with a continuous monoculture of crops may cause its
depletion to the point that soil amendment with soluble Si sources is needed to achieve high yields
(31). Some organic soils (histosols) contain little Si in their native state. Some, such as oxisols
and ultisols, are highly weathered, leached, and low in base saturation. Further, soils comprising
mainly quartz sand [silica (SiO2 )] (e.g., sandy entisols) may be high in insoluble Si but are very low
in plant-available Si. Other soil orders considered to be low or limiting are acidic alfisols, acidic
inceptisols, and spodosols (33, 159). Many soilless mixes fall into this category too. These low-
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soluble Si soil conditions are found in many crop-producing areas of Africa, Asia, the Americas,
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and Europe (33).

Soil weathering processes release SiO2 into the soil solution, surface water, and other natu-
ral water bodies mainly in the form of monosilicic acid (H4 SiO4 ), the prevalent and uncharged
monomeric molecule that exists below pH 9 (160). All plants grown in soil contain a certain con-
centration of Si in their tissues, as reported for more than 44 angiosperm clades that represent
over 100 orders or families (51, 71). When determining whether or not a plant accumulates Si,
past studies have focused primarily on measuring Si concentration in the foliage and not in other
plant organs (71, 154). However, coffee, peppers, and tomatoes concentrate as much, if not more,
Si in the roots as in the shoots (14, 52, 72). Si translocation from roots to shoots dramatically vary
among plant species and growth stage, and shoot Si concentration ranges from 0.1% to 10% on a
dry-weight basis (43). Monocots tend to accumulate more Si in shoots than do dicots. In general,
the Si shoot concentration is from 5% to 7% for wetland grasses, from 0.5% to 1.5% for dryland
grasses, and less than 0.5% for many dicots. At the low end of the range, 0.1% Si corresponds on
a dry-matter basis with macronutrients, such as Ca, magnesium (Mg), P, and S. At the upper end
of 10%, the tissue concentrations of Si exceed those of mineral nutrients such as nitrogen (N) and
K. Clearly, Si is a major mineral constituent of plants and is essential for diatoms, yellow-brown
or golden algae, and the Chrysophyceae and Equisetaceae, and is considered to be quasi-essential
for the growth and development of many plant species (43, 101, 120, 122).
Plants uptake Si from the soil solution as H4 SiO4 and subsequently transport it from roots
to shoots either passively or actively (101). The active mode of Si uptake from roots to shoots
was identified to be due to specific transporter proteins (10). These were first characterized in
certain monocots, such as barley, maize, rice, and wheat (21, 99, 100, 101, 110, 154), but dicots
such as cucumber, pumpkin, and soybean also have been found to have similar transporters (35).
Consequently, the difference in Si shoot concentration among plant species is attributed to their
differential ability to uptake this element by roots (10). Once across the endodermis, Si moves
through the xylem for plant species known to have the Si transporter proteins and/or is driven by
transpirational flow into the root endodermis, the cell membranes of the vascular bundle, and the
leaf cells of the epidermis to be further deposited beneath the cuticle (99, 100). The polymerization
of converting H4 SiO4 into insoluble silica (SiO2 nH2 O, a.k.a. opal, silica gel, or phytoliths) can
occur in the roots but is more prevalent beneath the cuticle and in the plant cell walls.
Plants supplied with Si counteract many types of abiotic (e.g., cold, drought, heat, salinity,
and mineral deficiency or toxicity) and biotic (e.g., plant diseases and herbivores and other pests)
stresses more efficiently (3, 32, 107, 130, 138). The benefits of Si were largely overlooked until the

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beginning of the twentieth century partly because of the elements abundance in nature and the
lack of visible symptoms of deficiency, and mainly because plant physiologists largely disregarded
the possible effects of this element from any meaningful experiments. However, in field and
greenhouse environments, plants are constantly exposed to different types of stresses and seem
to suffer more when grown in soils low or limiting in plant-available Si. An awareness of Si
deficiency in soils has become recognized since the 1980s as being a limiting factor in overall plant
development and crop yield (33). Although world conferences on Si in agriculture have been held
in the United States (1999), Japan (2002), Brazil (2005), South Africa (2008), China (2011), and
Sweden (2014), with the overarching theme of providing a forum in which the benefits of Si in
agriculture are discussed among plant scientists, agricultural managers, and companies producing
Si materials, the use of this element in agricultural production has been slow to materialize in many
countries. This review aims to provide the current status of the potential of Si in ameliorating
both abiotic and biotic stress in important economic crops.
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Metal Toxicity
Si is recognized for decreasing metal toxicity, and some mechanisms have been suggested to explain
this effect (3). The decrease in availability of phytotoxic metals in soil might be due to the increase
in pH and changes in the speciation (i.e., chemical and physical form) of metals in the soil solution
by the formation of silicate complexes (3). Sodium metasilicate increased soil pH and decreased
exchangeable lead (Pb), thus reducing soil Pb availability (87). Increases in soil pH and reduction in
bioavailable cadmium (Cd) also were reported as a result of applying amorphous silica (87% SiO2 )
(131). Cadmium in soil was also found to be in the form of specific adsorbed (bound to carbonates)
or iron (Fe)-manganese (Mn) oxidebound fractions in Si-amended soil (93). Exogenously applied
Si decreased chromium (Cr) availability in the soil by promoting precipitation-bound and organic
matterbound Cr fractions (36). Formation of hydroxyaluminum silicate is thought to be involved
in Si-mediated detoxification of aluminum (Al) in plants (70). Si-rich biochar was demonstrated
to reduce the amount of soil-exchangeable Al and prevent its migration to the plant (119). The
reduced availability and phytotoxicity of metals afforded by corn cob silica were related to the
formation of insoluble Pb silicate, as revealed by X-ray diffraction analysis, and to an increase in
soil pH, which further promoted binding of the heavy metals to carbonates, Fe and Mn oxides,
organic matter, and S in the soil (145). Taken collectively, these results suggest that pH and changes
in metal speciation are targets for Si in alleviating metal toxicity. However, Si-modulated changes
in the rhizosphere rather than in the bulk soil and the effect of Si sources and its concentration
need to be explored to better understand the effect of this element in the immobilization of metals
Plants exposed to abiotic stress (e.g., drought, metal, and saline stresses) show an increased
concentration of reactive oxygen species (ROS), such as superoxide radical (O2 ), hydroxyl radical
(OH ), and hydrogen peroxide (H2 O2 ), that damage biomolecules such as nucleic acids, photosyn-
thetic pigments, and proteins (3, 96, 130, 144). A decrease in ROS production and an enhancement
in the antioxidant system for Si-supplied plants may account for their increased tolerance to abiotic
stresses. Application of Si and Si nanoparticles (SiNP) decreased O2 , H2 O2 , and malondialde-
hyde (MDA) concentrations in maize leaves by increasing activities of ascorbate peroxidase (APX),
dehydroascorbate reductase (DHAR), or glutathione (GSH) reductase (GR) under arsenic (As)
stress. This finding demonstrated that rising ascorbate (AsA)-GSH cycle components are one of
the key mechanisms to counteract As (157). Increases in the activities of APX, catalase (CAT), Silicons Role in Abiotic and Biotic Plant Stresses 4.3

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peroxidase (POX), and superoxide dismutase (SOD) in roots and leaves of cotton plants also were
related to Si-mediated reductions in H2 O2 and MDA concentrations under zinc (Zn) stress (5). A
similar mechanism was found to be operating in the Si-afforded alleviation of boron, Cd, Cr, Fe,
Pb, and Mn toxicity in several plant species (3).
The compartmentalization of metals in roots and shoots is another Si-mediated tolerance
mechanism. In early studies, Si was demonstrated to alleviate Mn toxicity without reducing its
foliar concentration in barley plants. Mn was found to be more evenly distributed in Si-supplied
plants rather than being concentrated within necrotic spots (165). Si increased metal concentration
in roots and, consequently, lowered shoot concentration. Increased copper (Cu) adsorption was
observed at the outer thin layer (<100 A) root surface and immobilized in the central cylinder of
the root epidermis, and translocation was limited through an enhanced thickening of a Si-loaded
endodermis (79). In addition, Si-treated cucumber plants displayed less Mn in the symplast than
in the cell wall (136). However, Si-induced Mn sequestration in nonphotosynthetic tissues was
suggested as a mechanism of tolerance in maize (39).
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Metal toxicity has also been reported to be decreased by coprecipitation of Si with metals
in roots and leaves. In rice, lowered Cd, Cu, Pb, and Zn translocation from culm to leaf by Si
amendments was attributed to their coprecipitation with this element (59). Si was also proposed
to form precipitates with Zn as well as change Zn2+ into nonphytotoxic forms in the cell wall of
less bioactive tissues, especially in sclerenchyma of roots (60). Similarly, a hemicellulose-bound,
complexing Cd form of Si with net negative charge was suggested to inhibit Cd uptake by rice
root cells (97).
Some studies also have reported that chelation of flavonoid-phenolics or organic acids with
metals takes place in mitigation of heavy metal stress by Si. In maize, higher exudation of Al-
chelating catechin and quercetin, as well as malic acid, was observed in Si plus Al-exposed maize
compared to Al alone (80). Furthermore, Si increased the proportion of inorganic and organic
anion ligands such as aconitate that increased Cu sequestration into a less toxic form (79).
Recently, Si was reported to not affect the expression of two metal chelators [MsPCS1 (phy-
tochelatin synthase) and MsMT2 (metallothionein)], but it increased that of Fe-related genes
[MsIRT1 (Fe transporter), MsNramp1 (metal transporter), and OsFRO1 (ferric chelate reductase)]
under Cd stress in alfalfa roots. This finding suggested that limiting Fe uptake rather than metal
sequestration is important in Si-mediated alleviation of Cd toxicity (76). Genes, e.g., PCS1, SAP1,
and SAP14, encoding members of the stress-associated protein (SAP) family and the transcription
factor genes AP2/Erf020, Hsf31, and NAC6 were upregulated in roots of Cd-stressed rice plants
but downregulated in the presence of Si and indicated that post-stress addition of Si was able to
reverse Cd stress (44). A study of the rice leaf proteome under Cd stress showed that 50 proteins,
including those associated with photosynthesis, redox homeostasis, protein regulation/synthesis,
pathogen response, and chaperone activity, were upregulated by Si, whereas a class III POX and
a thaumatin-like protein were found to be upregulated by Si irrespective of Cd (113). Therefore,
Si appears to regulate gene expression to alleviate metal toxicity.

Drought and Salt Stress

Si has been shown to decrease drought stress in plants through different mechanisms, which in-
clude an increase in mineral nutrient uptake, modification of gas exchange attributes, osmotic
adjustment, reduction in oxidative stress, modification of gene expression, and regulation of com-
patible solutes and phytohormone synthesis (130). Si-induced increases in a balanced mineral
nutrient uptake and translocation may account for observed beneficial effects. In rice, Si mitigated
the adverse effects of drought while increasing K and total P concentrations in straw (41). Si uptake

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was strongly correlated with K under water deficit in Rhodes grass (Chloris gayana), Timothy grass
(Phleum pratense), Sudan grass (Sorghum sudanense), and tall fescue (Festuca arundinacea) (42). Sim-
ilarly, Si increased Ca, K, and Mg concentrations in tomato seedlings under salt stress improving
their tolerance to salinity (86).
Impairments in gas exchange attributes and leaf relative water content (RWC) are among the
primary mechanisms by which drought and salt stress reduce crop productivity. Si has been demon-
strated to nullify such stress-induced impairments. For example, photosynthesis, transpiration rate,
and leaf water content under water stress were preserved in leaves of tomato plants supplied with
Si, suggesting that increased water uptake/transport rather than decreased transpiration was be-
hind Si-alleviated stress (144). Conversely, a Si-induced increase in water-use efficiency (WUE) in
maize subjected to water stress was related to a reduced transpiration rate through stomata, indi-
cating that Si affected stomatal aperture (54), which might be species-dependent. The increases in
RWC or WUE due to increased silicification of leaves or root endodermis that prevent water from
escaping through plant tissues are thought to be involved in Si-alleviated osmotic stress (130). Si
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alleviates salinity stress in tomato seedlings by increasing photosynthesis, leaf transpiration rate,
and stomatal conductance and by decreasing the concentrations of Na+ and Cl in the leaves stem
and roots (86). Photosynthesis, stomatal conductance, transpiration rate, WUE, the number of
stomata, and stomatal size in okra plants exposed to salt stress were increased by foliarly applying
Si (150 mg/L) (1). Cucumber plants supplied with Si and exposed to salt stress displayed increased
photosynthesis, transpiration rate, leaf water content, and root hydraulic conductance, a fact that
was ascribed to a decrease in Na+ and an increase in K+ concentrations in the leaves (163). Si is
known to reduce Na+ uptake by stimulating the root plasma membrane H+ -ATPase activity (166)
and to reduce Na+ translocation by promoting its binding to cell walls, thus reducing Na+ in the
leaf apoplast (63).
Salt toxicity and water deficit are known to induce lipid peroxidation by ROS in plants, and
Si was able to alleviate salt- and drought-induced oxidative stress by stimulating the antioxidant
system (1, 86, 96, 144). Tomato plants supplied with Si and submitted to water stress displayed
increased CAT and SOD activities, high levels of AsA and GSH, lowered electrolyte leakage and
O2 and H2 O2 concentrations, and no changes in the plasma membrane integrity as compared
with control plants (144). The H2 O2 and lipid peroxidation levels from drought-stressed wheat
plants supplied with Si were reduced and the concentrations of AsA, GSH, total phenolics, and
total flavonoids increased (96). In salt-stressed roots of tomato seedlings, CAT and SOD activities
and the concentrations of nonprotein thiols and AsA were increased, whereas H2 O2 and membrane
oxidative damage were decreased by Si (86). The CAT, POD, and SOD activities increased and
lipid peroxidation was reduced for both a salt-tolerant and a salt-sensitive okra genotype supplied
with Si and subjected to NaCl stress (1).
Regulation of compatible solutes and phytohormone synthesis, including abscisic acid (ABA),
glycine betaine, indole-butyric acid (IAA), jasmonic acid ( JA), polyamines (PAs), proline, and sal-
icylic acid (SA) may enhance plant tolerance to salt or water stress (1, 168). Proline and glycine
betaine concentrations increased for pepper plants supplied with Si and exposed to drought stress.
Furthermore, Si increased nitrate reductase activity, indicating that this element can also in-
crease drought tolerance by enhancing N assimilation (116). Levels of soluble sugars and levels of
both free and conjugated PAs, including putrescine, spermidine, and spermine increased, whereas
1-aminocyclopropane-1-carboxylic acid (ACC), the precursor of ethylene (ET) decreased upon
Si treatment of drought-stressed sorghum plants (168). Similarly, Si application reversed the
decrease in the concentrations of the PAs putrescine and spermine in the roots of salt-stressed
cucumbers (163). The higher accumulation of glycine betaine, proline, total free amino acids, total
soluble sugars, and total phenolics under Si supply indicated its effect in improving the osmotic Silicons Role in Abiotic and Biotic Plant Stresses 4.5

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adjustment capacity and antioxidant activity of okra plants under saline conditions (1). Short-term
Si applications downregulated JA and upregulated ABA after 6 and 12 h in rice plants under salinity
stress (82). In soybean, Si alleviated the adverse effects of NaCl on plant growth by enhancing
endogenous gibberellic acid and decreasing ABA levels (85). In another study, both JA and salicylic
acid (SA) concentrations declined for soybean plants supplied with Si and submitted to drought
stress, whereas JA and SA concentrations decreased and increased, respectively, for Si-supplied
plants under salt stress (64).
Quantitative real-time polymerase chain reaction analysis revealed that Si application increased
the expression of four AsA-GSH cycle genes and five flavonoid biosynthesis pathway genes in
drought-stressed wheat plants and decreased MDA and H2 O2 concentrations, indicating that this
element played a pivotal role in the coordinated transcriptional regulation of multiple antioxidant
defenses in response to drought (96). In sorghum, the Si-mediated upregulation of several root
aquaporin genes was thought to increase root hydraulic conductance and improve root water up-
take under osmotic stress (95), whereas an upregulation of key PA synthesis genes was also believed
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to be involved in Si-mediated salt tolerance (169). The application of Si in combination with salt
stress transiently increased the expression of the ABA biosynthesis-related genes zeaxanthin epox-
idase and 9-cis-epoxicarotenoid oxygenase 1 and 4 (ZEP, NCED1, and NCED4) in comparison to
that with salt stress alone in rice (82). In tobacco BY-2 cell cultures, Si alleviated the damage of
NaCl stress by increasing ET emission and the expression of ET biosynthesis genes, whereas Si
failed to enhance plant tolerance in the absence of ET (89).

Other Abiotic Stresses

In addition to the stresses reported above, Si was also demonstrated to alleviate stresses caused
by low temperature (75, 91), ultraviolet-B (UV-B) radiation (158), and nutrient deficiency (106,
115). In cucumber plants grown under chilling temperatures (15/8 C), exogenous Si decreased
leaf withering, O2 , H2 O2 , and MDA concentrations through increased APX, GR, GSH POX,
monodehydroascorbate reductase, POX and SOD reductase activities, and the concentrations of
both AsA and GSH (75). In addition, a higher deposition of Si in leaves of chilling-stressed seedlings
played a role in constraining accumulation of MDA and damage to lipid membranes (75). Similarly,
Si-supplied plants from a freezing-susceptible and a freezing-tolerant wheat cultivar grown at
5 C displayed higher water retention, CAT, and SOD activities and higher concentrations of
AsA and GSH, which were related to lowered accumulation of H2 O2 and MDA (91). Recently,
SiNP was found to more effectively alleviate UV-B stress in wheat than Si by triggering the
antioxidant defense system. The reduced membrane lipid peroxidation and electrolyte leakage
and ROS-induced damage to photosynthesis were nitric-oxide-signaling dependent (158).
Regarding mineral nutrient deficiencies, ameliorative effects of Si have been consistently ob-
served for K (17, 106), P (98, 114), and Fe (9, 115), but inconsistent results were reported for
Mn and Zn (9). Addition of sodium silicate increased K concentration in leaves, stems, and roots
of soybean plants under K deficiency and ameliorated K deficiencyinduced reductions in shoot
and root growth. Moreover, Si prevented oxidative stress and lipid peroxidation due to K defi-
ciency by increasing CAT, POX, and SOD activities (106). Si alleviated the decreases in the leaf
gas exchange parameters, whole-plant hydraulic conductance, and root hydraulic conductance as
well as increased K accumulation in the xylem of K-deficient sorghum plants, indicating that Si
improved plant-water status under K deficiency (17). Si was beneficial for rice growth only under
low (0.014 mM) or high (0.70 mM) P levels, an effect that was ascribed to decreases in Fe and
Mn uptake at low P levels and to a reduced P uptake at high P levels (98). Soil P sorption was
reduced by sodium silicate, and Si was involved in the ameliorative effects for maize growth under

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P deficiency (114). Under Fe-deficient conditions, accumulation of Fe-mobilizing compounds and

expression levels of genes encoding proteins involved in Fe acquisition (FRO2, IRT1, and HA1) in
roots of cucumber were increased by Si (115). The alleviating Si effect in Fe deficiency in cucum-
ber was also linked to an enhanced Fe distribution toward apical shoot parts due to an increased
accumulation of Fe-mobilizing compounds such as citrate and catechin (9). These findings are
consistent with reports showing that Si can increase the accumulation of transcripts related to
both the shikimate and phenylpropanoid pathways (SK and PAL, respectively) in root tips under
Fe deprivation (115).


Although Si has proven to be effective in controlling a high number of insect herbivores and other
arthropods (90) (Table 1), the exact nature of Si-mediated antiherbivore defense remains largely
unknown (129). Even so, some mechanisms underlying Si-afforded plant resistance to pests are
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very similar to what is known about the role this element plays in suppressing plant diseases.
For example, the proposed Si deposition beneath the cuticle acting as a physical barrier against
plant disease appears to also function in defense against a number of herbivores and other pests
The exposure of Spodoptera exempta to high-Si diets reduced leaf digestibility and caused
mandible wear, which probably impacted herbivore performance (102). Similarly, the reduced
food quality and food conversion efficiencies were suggested to be responsible for a decrease in
the third instars weight gain and pupal weight of the rice leaf folder (Cnaphalocrocis medinalis) at a
high Si rate (65). In rice, lower relative growth rates and reduced boring success of the sugarcane
borer (Diatraea saccharalis) were observed in plants supplied with Si in comparison to controls
(147). High foliar Si concentration was associated with reduced sugarcane stalk larval penetration
by the pyralid borer Eldana saccharina (84). Energy dispersive X-ray microanalysis revealed that Si
concentrations increased in the stalks of sugarcane supplied with Si, but the magnitude of accumu-
lation was tissue dependent. For example, the epidermis of the internode and root band displayed
high Si accumulation, whereas Si was not evident in the bud scale. This finding might explain
why the leaf bud is a preferred entry point for the larval stage of E. saccharina (78). Furthermore,
irrespective of the Si supply, a higher Si accumulation was observed in a borer-resistant sugarcane
cultivar than in a borer-susceptible sugarcane cultivar, suggesting that resistance may be, at least
in part, associated with the plants ability to accumulate Si in the shoots (78).
The first evidence that Si might play an active role in host resistance against herbivores was
reported by McNaughton & Tarrants (104). The authors showed that heavily grazed African
grasses accumulated more Si in their leaf blades than did less heavily grazed plants, indicating
that silicification was probably an inducible defense against mammalian herbivores. In another
study, Si applied to the soil or foliage showed that the greenbug aphids (Schizaphis graminum)
stylet penetration into wheat was not affected, thus challenging the physical barrier hypothesis (58).
Furthermore, biochemical analysis confirmed that Si enhanced wheat defense mechanisms against
the greenbug aphid through an increase in POX, polyphenoloxidase (PPO), and phenylalanine
ammonia lyase (PAL) activities, especially when plants received Si before aphid infestation (57).
Ranger and his colleagues (123) observed reductions in total cumulative fecundity and the intrinsic
rate of increase of Myzus persicae on plants of Zinnia elegans supplied with Si. These reductions
were associated with great concentrations of 5-caffeoylquinic acid, p-coumaroylquinic acid, and
rutin in Z. elegans plants supplied with Si as compared to those that were not supplied with Si.
Rice plants grown in a Si-amended soil and infested with the leaf folder (Cnaphalocrocis medinalis)
displayed higher CAT, PAL, POX, PPO, and SOD activities and lower MDA concentration than Silicons Role in Abiotic and Biotic Plant Stresses 4.7

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Table 1 Herbivores and other pests reported to be suppressed by silicona

Mouthpart Pests
Chewing African armyworm (Spodoptera exempta)
African striped rice stem borer (Chilo zacconius)
Asiatic rice borer (Chilo suppressalis)
Central shoot fly (Atherigonaindica infuscate)
Green rice caterpillar (Naranga aenescens)
Hessian fly (Mayetiola destructor)
Hessian fly (Phytophaga destructor)
Leaf folder (Cnaphalocrocis medinalis)
Pink stalk borer (Sesamia spp.)
Sugarcane borer (Diatraea saccharina)
Stalk borer (Eldana saccharina)
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Sunflower caterpillar (Chlosyne lacinia saundersii )

Yellow rice borer (Scirpophaga incertulas)
Weevil (Cylas formicarius)
Piercing-sucking Aphid (Sitobion avenae)
Brown planthopper (Nilaparvata lugens)
Greenbug (Schizaphis graminum)
Green leafhopper (Nephotettix bipunctatus cincticeps)
Green peach aphid (Myzus persicae)
Small brown planthopper (Laodelphax striatellus)
Spider mite (Tetranychus urticae)
White-backed planthopper (Sogatella furcifera)
Whitefly (Bemisia tabaci )

Adapted from Reference 90.

plants grown in a nonamended soil. This finding suggested that Si-supplied plants were primed
to enhance antioxidant metabolism and defense mechanisms upon pest attack (66). The level of
transcripts for defense marker genes, JA accumulation, and POX and PPO activities, as well as
the level of trypsin protease inhibitor, increased in rice plants supplied with Si and attacked by
C. medinalis (167). In addition, JA was found to be important in the rice resistance against leaf
borer attack because Si was not able to increase resistance to this insect in plants whose genes
involved in JA biosynthesis or JA perception were silenced (167). Interestingly, Si deposition was
reduced when JA signaling was compromised, indicating a strong interaction between JA-mediated
responses to insect herbivores and Si as well as promotion of Si accumulation and JA (167).
Besides affecting bitrophic (plant-herbivore) interactions, Si has also been shown to affect
tritrophic (plant-herbivorenatural enemy) interactions, which may account for another possible
role in controlling pests (83, 129). In this case, Si may reduce insect feeding either by making them
more vulnerable to their natural enemies (55) or by changing the composition and concentration
of herbivore-induced plant volatiles (HIPVs), thus attracting the insects enemies (83). Although
potassium silicate applied at higher concentrations (0.08 and 0.16%; v/v) alone did not kill mites
(Tetranychus urticae), Beauveria bassianainduced mite mortality was enhanced for beans, cucum-
ber, eggplant, and maize (55). This finding was attributed to Si-primed biochemical defenses in
the plants, making the mites more susceptible to the entomopathogen (55). In rice, a combined

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application of potassium silicate, B. bassiana, and imidacloprid resulted in the maximum mortality
of C. medinalis (141). Cucumber plants supplied with Si were more attractive to adults of the preda-
tor Dicranolaius bellulus when attacked by larvae of Helicoverpa armigera in comparison to those
not supplied (83). The number of eggs from H. armigera removed from potted cucumber plants
and placed in a lucerne stand with numerous predatory arthropods was higher in the presence of
Si (83). A change in the HIPVs profile produced by Si-supplied cucumber plants when attacked
by the herbivore was thought to be involved in the plants attractiveness to natural enemies (83).
Indeed, considering that JA signaling is activated by Si for plants attacked by pests (167), such
a pathway could provide HIPVs required for the natural enemies attraction (129). Conversely,
even though the preference of the greenbug aphid (S. graminum) for wheat plants supplied with Si
was decreased, the attractiveness of the natural enemies Chrysoperla externa and Aphidius colemani
was not observed (111). Reynolds et al. (129) pointed out that effects of HIPVs might not have be-
come apparent because wheat plants were narrowly spaced and a nonchoice design was employed.
Therefore, Si seems to play an important role in the complex mechanisms involved in decreasing
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pests attack, which include physical barriers as well as biochemical and molecular changes that
can either directly reduce pest feeding and development or enhance the plants attractiveness to
natural enemies of pests.


A supply of Si to plants has been shown to reduce the intensities of several diseases [e.g., damping
off, leaf blights, leaf spots, galls, powdery mildews, root rots, rusts, and wilts; caused by different
genera of bacteria, fungi, nematodes, and oomycetes as well as a virus (Table 2)] in many eco-
nomically important crops (50, 132). In many of the host-pathogen interactions studied, both final
disease severity and the area under the disease progress curve were reduced as a result of the posi-
tive effect of Si on key components of host resistance, such as the incubation period, latent period,
colony or lesion size, lesion number, lesion expansion, and inoculum production per infection
site (50, 132). Sis impact on components of host resistance has helped to partially explain why

Table 2 Genera of plant pathogens causing seedborne, soilborne, and foliar diseases that had
their infection process impaired by silicona
Pathogens Genera
Fungi Alternaria, Bipolaris, Blumeria, Botrytis, Bremia, Cylindrocladium, Cercospora,
Colletotrichum, Corynespora, Diaporthe, Didymella, Drechslera, Erysiphe, Fusarium,
Ganoderma, Hemileia, Leptosphaeria, Magnaporthe, Microdochium, Monilinia,
Monographela, Mycosphaerella, Oculimacula, Oidiopsis, Oidium, Penicillium,
Pestalotia, Phakopsora, Phomopsis, Podosphaera, Pseudocercospora, Puccinia,
Pyricularia, Ramularia, Rhizoctonia, Sclerospora, Sclerotinia, Septoria, Sphaerotheca,
Trichothecium, Uncinula, and Ustilago
Oomycetes Phytophthora and Pythium
Bacteria Acidovorax, Pseudomonas, Ralstonia and Xanthomonas
Virus Nepovirus
Nematode Meloidogyne

A more comprehensive species list of fungi, oomycetes, bacteria and nematodes causing plant diseases that are suppressed
by silicon in a number of economically important crops can be found by consulting the Reference 132. Silicons Role in Abiotic and Biotic Plant Stresses 4.9

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the resistance of susceptible cultivars may be enhanced to the level of cultivars with race-specific
resistance (128, 134, 139).
The physical barrier hypothesis, which is related to Si deposition and polymerization beneath
the cuticle, in the cell wall, and inside the bulliform cells, was first proposed to explain the reduction
in rice blast severity (Figure 1a). Indeed, this hypothesis was also accepted for explaining the
reductions in disease intensities occurring in other monocots as well as in some dicots species.
In the case of the ricePyricularia oryzae interaction, fungal penetration can be either avoided
(reduction in the number of infection pegs formed by appressoria that pierced the underlying cell

Polymerized silicon Soluble silicon Necrotic cell

a ap
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Homogeneous Si deposition
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High cu

Low ep

No lesions me

Disuniform Si deposition



Small lesions

Reduced Si deposition



Large lesions

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wall; consequently, lesions did not develop) or delayed (increase in both incubation and latent
periods and decrease in lesion size) (2, 67, 69, 73, 135, 140, 153, 170). An outer, electron-dense
Si layer embedded in the cellulose microfibrils and an inner electron-translucent layer in the
epidermal cell wall of leaves of rice plants supplied with Si inhibited the colonization by P. oryzae
(81). The disuniformity in Si deposition beneath the cuticle and its further polymerization in the
form of amorphous silica bodies in the rice epidermal cells may allow the narrow penetration
peg of P. oryzae to penetrate (Figure 1b), but a delay in the incubation period and even in the
latent period can occur (12, 81, 112, 135). The reduced Si deposition beneath the cuticle and
the few epidermal cells containing silica bodies allowed penetration by P. oryzae. In this case,
both the incubation period and latent period were shorter, and the sizes of the blast lesions were
larger as a consequence of an unlimited colonization of the epidermal and mesophyll cells by
fungal hyphae (Figure 1c) (135). The silicification of oat epidermal cell walls reduced penetration
by Blumeria graminis f. sp. avenae (15). According to Seebold et al. (140), Si acts more than
as a physical barrier to increase rice resistance to blast because developing leaf lesions were of
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reduced size on rice cultivars with different basal levels of resistance. For the Arabidopsis thaliana
Erysiphe cichoracearum, cucumberColletotrichum lagenarium, cucumberPodosphaera xanthii, rice
Monographella albescens, riceRhizoctonia solani, wheatBipolaris sorokiniana, and wheatP. oryzae
interactions, the silicification of the epidermal cells delayed fungal penetration, colonization, and
sporulation (6, 30, 38, 46, 77, 105, 135, 137, 150). Appressoria of B. graminis f. sp. hordei were
unable to penetrate the epidermal cell walls of barley with high Si deposition. However, if fungal
penetration occurred, Si was deposited in the papillae and in the haustorial neck in an attempt to
delay penetration and haustoria formation (15, 74, 171). The deposition of Si along with phenolics
and hydroxyproline-rich glycoproteins in the leaves of French bean and cowpea plants reduced
the number of haustoria formed by Uromyces vignae (68, 117). Papillae formation seemed to be
stimulated in the presence of Si as reported for the rosePodosphaera pannosa and wheatB. graminis
f. sp. tritici interactions (8, 143). The interruption in Si supply resulted in high powdery mildew
and blast severities on the leaves of cucumber and rice plants, respectively. This finding suggested
that a minimal Si concentration and its continuous supply are required in newly formed leaves for

Figure 1
Proposed hypotheses to explain how silicon (Si) suppresses leaf blast, caused by Pyricularia oryzae,
development on rice. (a) The physical barrier hypothesis: The homogeneous Si deposition and
polymerization beneath the cuticle, as determined by X-ray microanalysis, completely block fungal
penetration, and, consequently, blast lesions do not develop. (b) The Si-mediated host defense mechanisms
hypothesis: The disuniformity in Si deposition beneath the cuticle, as determined by X-ray microanalysis,
may allow the narrow penetration peg to penetrate the cuticle and the wall of the epidermal cell. In this case,
there is a delay in both the incubation period (appearance of symptoms) and latent period (fungal
sporulation). Blast lesions are reduced in size owing to limited colonization of the leaf tissues by fungal
hyphae. A delay in fungal penetration allows the cells to counteract against fungal infection more efficiently
through the activation of host defense mechanisms (e.g., production of high amounts of phenolics,
phytoalexins, and lignin as well as high activities of defense enzymes such as chitinases and -1,3-
glucanases). It is believed that the soluble Si at the infection sites may be involved somehow in the
potentiation of these host mechanisms of defense. (c) Plants not supplied with Si: The reduced Si deposition
beneath the cuticle, as determined by X-ray microanalysis, allows fungal penetration. In this case, both the
incubation period and latent period are shorter, and the size of the blast lesion is larger as a consequence of
an unlimited colonization of the epidermal and mesophyll cells by fungal hyphae. In the X-ray microanalysis
images of the adaxial surface of rice leaves, red indicates the highest Si deposition and blue or black indicates
the absence of Si. Abbreviations: ap, appressorium; c, conidium; cu, cuticle; cw, cell wall; ep, epidermis; gt,
germ tube; me, mesophyll; sl, silica layer. Silicons Role in Abiotic and Biotic Plant Stresses 4.11

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optimal disease control (137, 152). Furthermore, it is postulated that the insoluble Si may not be as
important as the soluble Si at the time of fungal ingress to lower the disease intensity. The physical
barrier hypothesis is probably one tiny facet of how Si suppresses plant disease development, and
one must now take into account that the physiological, biochemical, and molecular data now
available will open new avenues of research to better understand the role of Si as a modulator of
host defense mechanisms.
Microscopic analyses showed that Si potentiated host defense mechanisms as evidenced by the
fact that many leaf cells of cucumber plants supplied with Si rapidly accumulated phenolics in
response to infection by Podosphaera xanthii (105). Biochemical analyses revealed high concen-
trations of flavonoids and phenolic acids in the leaf extracts of cucumber plants supplied with
Si, which also showed reduced powdery mildew symptoms (47). At both histopathological and
ultrastructural levels, many empty hyphae of P. oryzae and M. albescens were found to be evenly
surrounded or trapped in a dense osmiophilic material that also partially occluded epidermal and
mesophyll cells and the vascular bundle of rice plants supplied with Si (6, 135). In contrast to the
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Si-supplied plants, the epidermal and mesophyll cells of nonsupplied rice plants were devoid of
organelles, and the cell walls were no longer discernible in the mesophyll. In addition, the vas-
cular bundle was massively colonized by fungal hyphae (6, 135). Intense and rapid deposition of
electron-dense phenolic-like material was found in the root cells of cucumber plants supplied with
Si and infected with Pythium ultimum (20). Cytologically, leaves of wheat plants supplied with Si
reacted against infection by B. graminis f. sp. tritici as evidenced by an intense autofluorescence of
the epidermal cells (an indicator of phenolics) and increased concentration of fungitoxic aglycones
(125). The resistance of wheat plants supplied with Si to powdery mildew was linked to a high
concentration of antifungal methylated forms of trans-aconitate that possibly acted as phytoalexins
(126). Histochemically, Si proved to enhance the deposition of dopamine, flavonoids, lignin, and
phenolics in the sclerenchyma cells and in the metaxylem and phloem vessels in the roots of banana
plants infected with Fusarium oxysporum f. sp. cubense (48). Hyphae of F. oxysporum f. sp. vasinfectum
found in the endodermal cells and vascular vessels in the roots of cotton plants supplied with Si
were degraded due to the deposition of an electron-dense material (164). In the wheatP. oryzae
interaction, the production of phenolics and flavonoids was enhanced in the infected leaves of
Si-supplied plants (22, 30). Glycosylated phenolics, based on cytochemical labeling, were densely
deposited on the extrahaustorial membrane of the fungus and in the cell wall of epidermal cells of
wheat plants supplied with Si and infected with B. graminis f. sp. tritici (8).
In general, the activities of several enzymes related to host defense (e.g., -1,3-glucanase
(GLU), chitinase (CHI), PAL, POX, and PPO increased in plants supplied with Si as reported for
the following host-pathogen interactions: bananaF. oxysporum f. sp. cubense, bananaXanthomonas
campestris pv. musacearum, bitter gourdErysiphe sp., cucumberP. ultimum, cucumberP. xanthii,
coffeeMeloidogyne exigua, cottonColletotrichum gossypii var. cephalosporioides, cottonPhakopsora
gossypii, cottonRamularia areola, melonPodosphaera xanthii, peaMycosphaerella pinodes, perennial
ryegrassP. oryzae, riceB. oryzae, riceM. albescens, riceP. oryzae, riceR. solani, riceXanthomonas
oryzae pv. oryzae, sorghumColletotrichum sublineolum, soybeanPhakopsora pachyrhizi, and tomato
Pseudomonas syringae pv. tomato (4, 13, 19, 23, 25, 2729, 49, 61, 62, 92, 103, 121, 124, 128, 148, 149,
156, 172). Based on two-dimensional electrophoresis maps, Chen et al. (18) found that 26 proteins
were altered (16 increased and 10 decreased) in the roots of tomato plants supplied with Si and
infected with Ralstonia solanacearum, and most of them were associated with both plant metabolism
and defense mechanisms. A high Si concentration in the shoots of plants infected with different
pathogens was associated with a more efficient antioxidant metabolism (high APX, CAT, GR, and
SOD activities), thereby enhancing the removal of ROS (24, 34, 37, 49, 88, 108, 118, 127, 152).
The reduced ROS-dependent cellular damage was indirectly linked to lower MDA concentration

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due to reduced lipid peroxidation in Si-supplied and pathogen-infected plants (24, 28, 34, 37, 49,
109, 118, 127). Debona et al. (34) reported that wheat plants supplied with Si and infected with
P. oryzae showed lower cellular damage and decreased APX, CAT, GSH-S-transferase, POX, and
SOD activities. Activities of APX, CAT, and SOD decreased in the leaves of wheat plants supplied
with Si and infected with B. graminis f. sp. tritici (109). From a proteomic perspective, Liu et al.
(94) reported that the quantity of APX, DHAR, and SOD increased in rice plants supplied with
Si and infected with P. oryzae.
Si has been shown to potentiate host defenses by enhancing defense-related gene expres-
sion. The genes encoding for GLU, chalcone synthase, CHI, cinnamyl alcohol dehydrogenase,
lipoxygenase, PAL, POX, and PR-1 were all upregulated in the perennial ryegrassP. oryzae,
riceP. oryzae, rosePodosphaera pannosa, and wheatP. oryzae interactions (22, 121, 135, 142).
For snap beans, genes encoding an ET response factor, a polygalacturonase inhibitor protein,
a defense-associated phosphatase, and PR-1 were overexpressed and contributed to resistance
against gray mold (Botrytis cinerea) and cottony rot (Pythium aphanidermatum) (40). The expres-
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sion levels of the genes Os03g0109600 (encoding a protein with 175 amino acids and containing
a nuclear localization signal), Prla (PR-1), Rcht2 (chitinase), Lox2 (lipoxygenase), and Pal (phenyl-
alanine ammonia lyase) were high for rice plants supplied with Si and infected with X. oryzae pv.
oryzae, whereas the genes CatA (catalase) and Os03g0126000 (encoding a binding protein with 424
amino acids) had their expression downregulated (149). Many antimicrobial compounds, related
to both primary and secondary metabolism, such as anthocyanins, flavonoids, lignin-thioglycolic
acid derivatives, momilactones, and phenolics, had their concentrations increased in the tissues of
pathogen-infected plants supplied with Si (25, 28, 49, 61, 62, 121, 128, 135, 142, 148, 149, 164,
Some transcriptomic studies have been carried out to elucidate the molecular mechanisms
by which Si potentiates host defense. Fauteux et al. (45) used a 44-K microarray chip to study
the expression of genes of Arabidopsis plants nonsupplied or supplied with Si and infected with
B. graminis f. sp. tritici. The expression of all but two genes was unaffected by Si in noninoculated
plants. Regardless of Si, the expression of a set of nearly 4,000 genes was either upregulated [e.g.,
R genes, stress-related transcription factors, genes involved in signal transduction, the biosynthesis
of stress hormones (SA, JA, and ET) and the metabolism of ROS] or downregulated (genes involved
in primary metabolism) by pathogen infection. The magnitude of downregulation was dramatically
attenuated for the inoculated plants supplied with Si, indicating alleviation of the stress imposed
by the fungus (45). In another study, Chain et al. (16) performed transcriptomic analysis of 55,000
unigenes to compare the response of wheat plants nonsupplied or supplied with Si in the absence
or presence of infection by B. graminis f. sp. tritici. Only 47 genes of diverse functions and with
little evidence of regulation of a specific metabolic process were upregulated for the uninoculated
and Si-supplied plants. Many genes linked to stress and metabolic processes were upregulated, and
genes associated with photosynthesis were downregulated for infected plants not treated with Si.
However, for the infected and Si-supplied plants, genes linked to stress and metabolic processes
were downregulated, and those linked to photosynthesis were upregulated. The expression of
genes associated with JA/ET pathways, oxidative stress, and host defense had their expression
upregulated in the roots of tomato plants supplied with Si and infected with Ralstonia solanacearum
Brunings and her colleagues (11) used a 44-K gene chip to study gene expression of rice
plants treated with Si and nontreated and then infected by P. oryzae. They found that a
unique number of defense/stress-related genes [ET signaling pathway genes, a gene encoding
a thaumatin/pathogenesis-related protein (Os12g0568900), a class III POX (Os07g0677500) and
a number of transcription factors and protein kinases] were upregulated in the P. oryzae versus Silicons Role in Abiotic and Biotic Plant Stresses 4.13

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Si + P. oryzae treatment comparisons, highlighting how Si may have changed the plants reaction
to the pathogen. Further, the Si versus Si + P. oryzae comparison had 440 differentially expressed
genes fewer than the control versus P. oryzae comparison, and this comparison had only 236 genes
in common. Si therefore affected the interaction between the host and pathogen at the molecular
level by attenuating the rice response to the pathogen and by the differential expression of a unique
set of genes. Consequently, rice plants supplied with Si were preconditioned to react against this
pathogen stress, and this is further supported by the finding that infection by P. oryzae resulted
in less than half the number of differentially expressed genes in Si-amended plants versus nona-
mended plants (298 versus 738, respectively). All these aforementioned studies were unanimous in
concluding that Si played a pivotal role in the plants transcriptome only upon pathogen infection.
The physiology of plants infected by pathogens is negatively affected as evidenced by changes
in the values of leaf gas exchange parameters, reductions in the amount of healthy leaf area, de-
creases in the concentration of photosynthetic pigments, damage to the chloroplasts, impairments
in energy dissipation via chlorophyll a fluorescence and high foliar temperature (146). Several
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studies showed that Si partially preserved the photosynthetic performance of plants infected by
pathogens. For many host-pathogen interactions such as common beanColletotrichum lindemuthi-
anum, cottonR. areola, cottonC. gossypii var. cephalosporioides, riceP. oryzae, riceM. albescens,
sorghumC. sublineolum, and wheatP. oryzae, the Si supply to plants, besides reducing disease
severities, contributed to the preservation of photosynthesis and the concentration of photosyn-
thetic pigments (7, 24, 37, 61, 118, 127, 155). Some photochemical parameters associated with
chlorophyll a fluorescence, such as the quantum yield of photosystem II (PSII) photochemistry
(Fv /Fm ), the photochemical quenching coefficient, and the electron transport rate, were greatly im-
proved for rice plants infected with P. oryzae or M. albescens and wheat plants infected with P. oryzae
if supplied with Si (7, 37, 155). The heat dissipation of chlorophyll excitation energy, based on the
nonphotochemical quenching parameter, decreased for rice and wheat plants infected with either
P. oryzae or M. albescens, respectively, when supplied with Si compared to infected controls. This
finding suggests that the PSII electron transport at the chloroplast level was not impaired, and the
photoprotective processes were preserved by Si treatment (7, 37, 155). Gao et al. (53) also reported
that an increase in rice resistance to blast due to Si was associated with an enhancement in the
photochemical efficiency, specifically by an increase in the values of Fv /Fm and the maximum pri-
mary yield of the photochemistry of PSII. Bipolaris oryzae infection repressed photosynthesis and
nitrate reduction in the leaves of rice plants nonsupplied with Si, triggering premature senescence
and therefore contributing to an increase in brown spot severity (162). By contrast, the expression
of many photorespiratory marker genes was upregulated in Si-supplied plants, indicating that
an increase in photorespiration helped to alleviate the stress imposed by B. oryzae infection by
protecting the rices photosynthetic machinery (162). Several genes involved in photosynthesis
and metabolism of amino acids, carbohydrates, and lipids were downregulated in the leaves of
Arabidopsis thaliana nonsupplied with Si and infected by E. cichoracearum. However, in infected
plants supplied with Si, many of these genes were less responsive (45). For the wheatB. graminis
f. sp. tritici interaction, the supply of Si to plants contributed to the downregulation of genes
associated with stress and metabolic processes and the upregulation of those related to photo-
synthesis (16). Proteins involved in photosynthesis (chlorophyll a/bbinding protein, chloroplast
putative thylakoid lumenal 16.5-kDa protein, sedoheptulose-1,7-bisphosphatase, and ribulose bis-
phosphate carboxylase large subunit) and the tricarboxylic acid and pentose phosphate pathways
were upregulated in rice plants supplied with Si and infected with P. oryzae compared to in-
fected controls without Si (94). According to Dallagnol et al. (26), the inefficiency of the lsi1
(low Si; defective in active Si uptake) rice mutant, which originated from the wild-type cultivar
Oochikara, to actively uptake Si resulted in greater brown spot severity and, consequently, reduced

4.14 Debona Rodrigues Datnoff

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photosynthesis and foliar sugar concentration. By contrast, the high foliar Si concentration de-
creased brown spot symptoms, preserved photosynthesis, and promoted high foliar sugar concen-
tration for the wild-type cultivar Oochikara. Rice resistance to brown spot was independently and
additively affected by Si and soluble sugar concentrations in the leaf tissue (26).
Some studies have linked Si with the plant hormones that modulate defense mechanisms such
as ET, JA, and SA in the ArabidopsisE. cichoracearum interaction (45), ET in the riceP. oryzae
and riceB. oryzae interactions (11, 161), and ET and JA in the tomatoR. solanacearum interaction
(56). Rice resistance to brown spot potentiated by Si was associated with the inability of the ET
produced by B. oryzae to suppress the rice plants innate immune system and, consequently, to
avoid the activation of the basal mechanisms of defense (161).

Si, a major component of the soil-plant system, plays significant roles in a plants life especially in
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alleviating both abiotic and biotic stresses. Unfortunately, the recognition of this quasi-essential
element as one component of fertilizers that could be used in an integrated disease(s)/pest(s)
management strategy is still in its infancy. In general, plants having a high shoot or root Si
concentration are less prone to suffer from pest attack and exhibit enhanced tolerance to abiotic
stresses such as drought, metal toxicity, salinity, low temperature, and UV-B radiation. However,
the most notable effect of Si is the reduction in the intensities of a number of plant diseases caused by
biotrophic, hemibiotrophic, and necrotrophic plant pathogens causing seedborne, soilborne, and
foliar diseases in many crops of great economic importance. The reduction in symptom expression
can be ascribed to the effect of Si on components of host resistance, including incubation period,
latent period, lesion size, lesion number, and inoculum production per infection site. Although
not discussed in this review, the intensities of some diseases for Si-supplied plants are lowered to
the same level achieved with fungicides, and the resistance of susceptible cultivars amended with
Si may achieve a comparable level exhibited by cultivars with race-specific resistance (132).
The polymerization of Si beneath the cuticle and in the wall of epidermal cells is one hypothesis
that explains how this element reduces lesion number and size for some diseases. However, new
insights have revealed that in many plant species supplied with Si, the phenylpropanoid and
terpenoid pathways are greatly potentiated, and there is a faster and stronger transcription of
genes involved in host defense and higher activities of defense enzymes. Antioxidant metabolism
involved in the removal of ROS and photosynthesis were greatly improved in Si-supplied plants
regardless of the type of stress. Although our understanding of how this overlooked element helps
plants counteract pathogen infection and pest attack as well as different types of abiotic stress has
advanced, the exact mechanism(s) by which Si modulates plant physiology through the potentiation
of host defense mechanisms still needs to be investigated in more detail at the genomic, proteomic,
and metabolomic levels.

1. A minimum Si concentration in the roots and/or shoots of higher plants, especially mono-
cots, is needed to effectively slow disease development and pest attack and to alleviate
abiotic stresses (e.g., drought, low temperature, metal toxicity, salinity, and UV-B radia-
tion). As the Si concentration (insoluble or soluble) in plant tissue increases, suppression
of both abiotic and biotic stresses occurs more efficiently. However, Si needs to be made
available for plant uptake during their entire life cycle to reduce these stresses. Silicons Role in Abiotic and Biotic Plant Stresses 4.15

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PY55CH04-Datnoff ARI 8 May 2017 16:51

2. Many components of host resistance (e.g., incubation period, latent period, lesion num-
ber, lesion size, and amount of inoculum produced per infection site) are greatly affected
by Si contributing to reduce the apparent infection rate of a certain disease.
3. In the presence of Si, the resistance of some cultivars known to be susceptible may increase
to the same level of resistance of cultivars exhibiting race-specific resistance.
4. The response of plants against pathogen infection, pest attack, and even abiotic stresses
at the biochemical, physiological, and molecular levels are remarkably similar when Si
is taken up by roots and translocated to shoots, indicating an active role played by this
element in one or more plant defense signaling pathways.

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The authors are not aware of any affiliations, memberships, funding, or financial holdings that
might be perceived as affecting the objectivity of this review.

We acknowledge the financial support obtained from CAPES, CNPq, and FAPEMIG to sup-
port the studies investigating the effect of silicon to control plant diseases. Professor Fabrcio A.
Rodrigues thanks CNPq for his fellowship. Professor Lawrence E. Datnoff thanks the Louisiana
State University Agricultural Center for their generous operational and financial support.

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