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Annals of Applied Biology ISSN 0003-4746

RESEARCH ARTICLE

Silicon-augmented resistance of plants to herbivorous insects:


a review
O.L. Reynolds1 , M.G. Keeping2 & J.H. Meyer3
1 EH Graham Centre for Agricultural Innovation (New South Wales Department of Primary Industries and Charles Sturt University), Private Mail Bag,
Wagga Wagga, NSW 2650, Australia
2 South African Sugarcane Research Institute, Mount Edgecombe 4300, South Africa and School of Biological and Conservation Sciences, University
of KwaZulu-Natal, Scottsville 3209, South Africa
3 Agricultural Consultant, 16 Delaware Avenue, Durban North 4051, South Africa

Keywords Abstract
insect–plant interactions; trophic interactions;
resistance; induced plant defences; Silicon (Si) is one of the most abundant elements in the earth’s crust,
constitutive defences; biological control. although its essentiality in plant growth is not clearly established. However,
the importance of Si as an element that is particularly beneficial for plants
Correspondence
under a range of abiotic and biotic stresses is now beyond doubt. This paper
O.L. Reynolds (née Kvedaras), EH Graham
reviews progress in exploring the benefits at two- and three-trophic levels and
Centre for Agricultural Innovation (New South
Wales Department of Primary Industries and
the underlying mechanism of Si in enhancing the resistance of host plants to
Charles Sturt University), Private Mail Bag herbivorous insects. Numerous studies have shown an enhanced resistance of
Wagga Wagga, NSW 2650, Australia. plants to insect herbivores including folivores, borers, and phloem and xylem
Email: olivia.reynolds@industry.nsw.gov.au feeders. Silicon may act directly on insect herbivores leading to a reduction
in insect performance and plant damage. Various indirect effects may also
Received: 18 February 2009; revised version be caused, for example, by delaying herbivore establishment and thus an
accepted: 6 June 2009.
increased chance of exposure to natural enemies, adverse weather events or
control measures that target exposed insects. A further indirect effect of Si may
doi:10.1111/j.1744-7348.2009.00348.x
be to increase tolerance of plants to abiotic stresses, notably water stress, which
can in turn lead to a reduction in insect numbers and plant damage. There
are two mechanisms by which Si is likely to increase resistance to herbivore
feeding. Increased physical resistance (constitutive), based on solid amorphous
silica, has long been considered the major mechanism of Si-mediated defences
of plants, although there is recent evidence for induced physical defence.
Physical resistance involves reduced digestibility and/or increased hardness and
abrasiveness of plant tissues because of silica deposition, mainly as opaline
phytoliths, in various tissues, including epidermal silica cells. Further, there
is now evidence that soluble Si is involved in induced chemical defences to
insect herbivore attack through the enhanced production of defensive enzymes
or possibly the enhanced release of plant volatiles. However, only two studies
have tested for the effect of Si on an insect herbivore and third trophic level
effects on the herbivore’s predators and parasitoids. One study showed no
effect of Si on natural enemies, but the methods used were not favourable for
the detection of semiochemical-mediated effects. Work recently commenced
in Australia is methodologically and conceptually more advanced and an effect
of Si on the plants’ ability to generate an induced response by acting at the
third trophic level was observed. This paper provides the first overview of Si
in insect herbivore resistance studies, and highlights novel, recent hypotheses
and findings in this area of research. Finally, we make suggestions for future
research efforts in the use of Si to enhance plant resistance to insect herbivores.

Ann Appl Biol 155 (2009) 171–186 © 2009 The Authors 171
Journal compilation © 2009 Association of Applied Biologists
Silicon and insect herbivores O.L. Reynolds et al.

Introduction three ways. Firstly, bands of Si bodies may protect the


underlying vascular tissue restricting chewing or rasping
Silicon (Si) is only second in abundance to oxygen in the
herbivores to intercostal zones. Secondly, the silicifica-
earth’s crust, although its essentiality in plant growth is
tion of epidermal cell walls may resist entry in these
not clearly established. However, the importance of Si
areas. Thirdly, chewing herbivores may be inhibited from
as a nutrient that is particularly beneficial for plants
penetration by silicification of the leaf margins. Although
under a range of abiotic and biotic stresses, is now
evidence that supports the role of Si in the physical
beyond doubt. Insect herbivores represent one class
defence of plants against arthropods continues to grow, it
of biotic stressors that Si can provide some defence
was not until the work of Bélanger et al. (1995) and Fawe
against. Silicon is taken up by plants in the form of
et al. (1998), that the role of Si in amplifying the chemical
monosilicic acid and is transported from the roots to
defences of plants first became apparent. Fauteux et al.,
the shoots, and when concentrated over a critical level
(2005) proposed that Si may play two central roles in plant
(approximately 100 ppm at biological pH), it polymerises
chemical defence: (a) enhanced signal transduction at the
as opaline phytoliths, which comprise the bulk of a
single cell level leading to an increase in induced systemic
plant’s Si content (Jones & Handreck, 1967). Whether
resistance and (b) modulation of the generation of sys-
Si is taken up through the leaves remains controversial
temic signals, as both processes rely on primary elicitation.
(Guével et al., 2007). Although Si deposition in the Silicon may be applied as a post-harvest treatment
epidermal cells of plants is purportedly responsible for the against a broad range of agricultural and environmental
protective effects of Si against plant diseases and insect pests. Such treatments include novel silicon-containing
herbivores (Ma, 2004), there is increasing evidence for insecticides with activity against various species of termite
Si-induced chemical defence against plant diseases (Ma, and wood boring beetle in wood products (Adams
2004; Hammerschmidt, 2005), but only one published et al., 1995) and diatomaceous earth (a dust powder
example against insects (Gomes et al., 2005). formulation with silicon oxide) used as a dust toxicant
Silicon may act either directly or indirectly on insect against stored-grain insects (El-Lakwah et al., 2001; Lord,
herbivores (Kvedaras & Keeping, 2007). Direct effects 2001). In this paper, however, we review the role of Si
may include a reduction in insect growth and repro- as a pre-harvest treatment against herbivorous insects, at
duction with concomitant reduction in damage to the two and three-trophic levels and the possible defence
crop. Numerous studies have shown enhanced resistance mechanisms involved, addressing only those studies
of host plants treated with Si to insect herbivores and that have applied Si either via the roots or as a leaf
other arthropods, mostly at two-trophic levels (Djamin surface (foliar) application. Although the effect of Si on
& Pathak, 1967; Moraes et al., 2004, 2005; Kvedaras plant growth is particularly obvious under conditions
et al., 2007a,b, 2009; Kvedaras & Keeping, 2007). Indirect of stress, it is increasingly apparent that Si is involved
effects are those affecting insect mortality rates and may in complex defence mechanisms, most of which remain
result from delayed or reduced plant penetration, result- to be fully elucidated. Although several comprehensive
ing in increased exposure to natural enemies, adverse reviews have covered the role of Si in disease resistance,
climatic conditions and control measures that target there have been no such reviews of the role of Si in insect
exposed insects, such as chemical applications (Kvedaras herbivore resistance in the peer-reviewed literature. This
& Keeping, 2007). Indirect effects of Si may also occur paper provides the first overview of such studies, while
through increased tolerance of plants to abiotic stresses, highlighting novel, recent hypotheses and findings.
for example water stress, thus resulting in enhanced plant
resistance to insect attack (Kvedaras et al., 2007a). Recent
studies in Australia have also highlighted the potential for Two-trophic level studies
Si to act indirectly at the third trophic level by increasing
Silicon sources and insect resistance
the attraction of the natural enemies of an herbivorous
insect (Kvedaras et al., 2008; Kvedaras et al., 2009a). Various sources (or carriers; see Savant et al., 1999) of Si
Naturally occurring plant Si is a vital area to recognise have been used to deliver Si to the plant and test the
as the discovery of this association has led the way for efficacy of Si in enhancing host plant resistance to insects.
applied Si amendments to enhance plant resistance to These sources essentially fall into two classes: solid sources
insect herbivores. O’Reagain & Mentis (1989) proposed incorporated into the soil and silicate solutions applied as
that leaf silicification in grasses evolved in response to a soil drench or as a foliar spray.
invertebrate herbivory and that it is primarily aimed at Calcium silicate (Ca2 SiO4 ) (as an industrial slag by-
reducing tissue loss. They suggested that the histological product and from geological sources such as wollastonite
distribution of Si may inhibit small herbivore attack in (calcium meta-silicate; CaSiO3 )) has been a frequently

172 Ann Appl Biol 155 (2009) 171–186 © 2009 The Authors
Journal compilation © 2009 Association of Applied Biologists
O.L. Reynolds et al. Silicon and insect herbivores

chosen source for direct incorporation into the soil although Slagment® at a rate of 10 t ha−1 released
in experimental interaction studies between plant Si more than three times as much Si into the soil than
fertilisation and a range of insect herbivores including the USA calcium silicate, this greater release was not
stem borers (Anderson & Sosa, 2001; Keeping & reflected as increased leaf and stalk Si content, or greater
Meyer, 2002; Kvedaras et al., 2007b), phloem feeders reduction in borer performance and damage compared
(Correa et al., 2005; Goussain et al., 2005), and folivores with calcium silicate. Such studies demonstrate that
(Korndörfer et al., 2004; Redmond & Potter, 2007). In all although some sources provide large amounts of Si, much
cases, except the two studies cited above on folivores, of it may be unavailable to the plant, possibly because
the calcium silicate applications produced significant of re-polymerisation of silicic acid at high concentrations
reductions in insect performance and plant damage. following its initial solubilisation in the soil (Keeping &
Where silicate applications had no effect on insect Meyer, 2006).
performance and damage, they nevertheless elevated Sodium silicate (Na2 SiO3 ) solution has been an
the leaf Si content (Korndörfer et al., 2004; Redmond effective liquid source of Si in numerous Si/insect
& Potter, 2007). Among these studies, the amount interaction studies, either applied to the growth medium
of plant available Si in the calcium silicate varied as a soil drench (Basagli et al., 2003; Moraes et al.,
between 10% and 39%, with the product supplied 2004), or as foliar sprays alone (1, 2, and 4%;
at rates between 1.12 and 10.0 t ha−1 (Gomes et al., Hanisch, 1981) or as foliar sprays (0.5% or 1% SiO2 )
2005; Keeping & Meyer, 2006; Redmond & Potter, in combination with the soil drench (Moraes et al.,
2007). Calcium silicate has also been applied in aqueous 2004, 2005). Significant reductions in insect performance
solution as a foliar spray against whitefly, Bemisia tabaci were obtained, confirming that the Si source had
(Gennadius) (Homoptera: Aleyrodidae) (1.0% solution an effect (direct and/or indirect) on the target pest.
of ∼29% Si; Correa et al., 2005) and thrips, Thrips palmi For example, green bug Schizaphis graminum (Rondani)
Karny (Thysanoptera: Thripidae) (1.5% solution of ∼8% (Homiptera:Aphididae) reared on sodium-silicate-treated
Si; Almeida et al., 2008), with significant reductions in sorghum plants showed a reduction in feeding preference
insect performance (Correa et al., 2005; Almeida et al., and reproduction (Carvalho et al., 1999). Although
2008) and plant damage (Almeida et al., 2008). For application of sodium silicate and aphid pre-infestation
sugarcane/insect studies, bagasse furnace ash (or flyash, did not influence mortality and duration of the pre-
a common by-product of sugar mills produced from reproductive and reproductive period of Schiz. graminum
complete burning of bagasse) has also been used as on sorghum plants, fewer aphids chose leaf sections of
a Si source for direct incorporation into the soil (Pan plants subject to aphid pre-infestation or treated with
et al., 1979; Keeping & Meyer, 2006). Fly ash is an sodium silicate (Costa & Moraes, 2002), and aphid
inorganic source of calcium silicate, which includes colonisation of sodium-silicate-treated sorghum plants
smaller quantities of potassium silicate, sodium silicate was also reduced (Moraes & Carvalho, 2002). In wheat,
and magnesium silicate, as well as about 40% carbon sodium silicate application reduced Schiz. graminum
(dry weight). Porous hydrate calcium silicate applied to longevity and nymph production, as well as plant
turf grass (hybrid type of Zoysia japonica Steud. and Zoysia preference (Basagli et al., 2003). Italian ryegrass (Lolium
matrella (L.) Merr., cv. Miyako) produced a significant multiflorum L., cv. RvP) fertilised with sodium silicate
increase in wear resistance of the turf and a 41% decrease displayed a significant reduction in colonisation by the
in feeding by lawn cutworm, Rusidrina depravata Butler dipterous stem-boring larvae of Oscinella frit L. (Diptera:
(Lepidoptera: Noctuidae) (Saigusa et al., 1999). Keeping Chloropidae) and other related species (Moore, 1984).
& Meyer (2006) compared the effects of four Si sources To our knowledge, potassium silicate (K2 SiO3 ) has
incorporated in solid form, on Si uptake and resistance been used as a soil drench in only two published
of sugarcane to Eldana saccharina Walker (Lepidoptera: studies investigating the effects of applied Si on insect
Pyralidae); calcium silicate from the USA (12% pure herbivores: one on stem borer, Scirpophaga (Tryporyza)
Si; supplied by Calcium Silicate Corp®, Lake Harbour, incertulas (Walker) (Lepidoptera: Pyralidae) in rice
FL); wollastonite (12% Si; calcium meta-silicate, a by- (Subbarao & Perraju, 1976) and the other on leaf
product of the ceramics industry); Slagment® (18% Si; miner, Liriomyza trifolii (Burgess) (Diptera: Agromyzidae)
a blast-furnace slag supplied by Slagment Ltd., Alberton, in chrysanthemums (Parrella et al., 2007). In both,
South Africa), and flyash (10% Si). Silicon source had significant reductions in insect performance and increased
a large impact on both plant uptake of Si and on plant uptake of Si were recorded (Subbarao & Perraju,
the effect of Si on E. saccharina. For example, flyash at 1976; Parrella et al., 2007).
30 t ha−1 was less effective than calcium silicate at 10 t The effectiveness of foliar spray- versus root-applied
ha−1 in reducing E. saccharina infestation. Furthermore, soil drenches is discussed further below.

Ann Appl Biol 155 (2009) 171–186 © 2009 The Authors 173
Journal compilation © 2009 Association of Applied Biologists
Silicon and insect herbivores O.L. Reynolds et al.

Historical background to the role of Si in insect striatellus Fallen (Hemiptera: Delphacidae) (Liu et al.,
resistance 2007), Sitobion avenae (F.) (Hemiptera: Aphididae), and
Metopolophium dirhodum (Walker) (Hemiptera: Aphididae)
McColloch and Salmon (1923) were the first to sug-
(Hanisch, 1981), some xylem-feeding insects (e.g. Nila-
gest that silica played a role in the resistance of maize to
parvata lugens Stål. (Homoptera: Delphacidae)) (Yoshihara
hessian fly, Mayetiola destructor (Say) (Diptera: Cecidomyi-
& Sogawa, 1979), and other plant feeders (e.g. Cylas
idae). Later, Ponnaiya (1951) indicated that resistance
formicarius Fab. (Coleoptera: Curculionidae) (Singh et al.,
in sorghum to central shoot fly, Atherigona indica infus-
1993). In a later study, Massey et al., (2007) found that
cata Emden (Diptera: Muscidae) was related to Si. The
the phloem feeder, Si. avenae, suffered no detrimental
first study that showed an increase in plant resistance
effects from increased plant Si.
to an insect as a result of Si application was that on
rice stem borer, Chilo simplex Butler (Lepidoptera: Pyral-
Nutritional and physical defences
idae) (Sasamoto, 1953). Slag applied to rice plants in
Japan reduced Ch. simplex damage and increased growth The most widely accepted mechanisms for the action
impetus of the plant (Sasamoto, 1953), likely because of of Si in increasing plant resistance to insect attack
increased strength of the rice stem following Si accu- are reduced digestibility and increased hardness and
mulation (Sasamoto, 1955). Thereafter, studies have abrasiveness of (especially epidermal) plant tissues
increasingly shown enhanced plant resistance to insect because of silica deposition, mainly as opaline phytoliths,
herbivores in a range of crops fertilised with Si, with and in association with cell walls (Kaufman et al., 1985;
a growing focus on determining the underlying mech- Salim & Saxena, 1992; Panda & Khush, 1995; Ma et al.,
anism/s. More recently, Nakata et al., (2008) elegantly 2001; Massey et al., 2006; Massey & Hartley, 2009). These
demonstrated that Si accumulation by rice is involved in may affect the herbivore either directly or indirectly.
resistance to pests. They found that chewing damage from
rice leaf roller, Cnaphalocrocis medinalis (Guenée) (Lep- Nutrition
idoptera: Pyralidae) and rice green caterpillar, Naranga
Elevated levels of Si may increase the bulk density of
aenescens Moore (Lepidoptera: Noctuidae) was apparent
the diet such that insects are unable to ingest sufficient
in leaves of the low silicon rice 1 (lsil) mutant cloned
quantities of nutrients and water (Panda & Kush, 1995).
by Ma et al., (2006), which controls Si accumulation in
For example, Massey et al., (2006) showed that Sp. exempta
rice, but not in the wild type plant. Thus the mutant
fed on high-Si plants of three (out of five) grass species not
cannot acquire resistance to biotic stressors even under
only suffered reduced digestion efficiency, but increased
field conditions in which Si is at a high level.
its rate of consumption on two (of the five) grass
species only. Possibly, more specialist feeders such as
Sp. exempta have limited ability to increase consumption
Insect feeding guilds
to compensate for feeding on poorer quality host plants
Massey et al., (2006) postulated that plant Si was effective (Lee et al., 2003). Massey & Hartley (2009) subsequently
in defending against folivorous but not phloem-feeding found that Si reduced both the efficiency with which
insects. Such a differential effect of silicon between insect Sp. exempta converted ingested food into body mass
feeding guilds was contested by Keeping & Kvedaras and the amount of nitrogen absorbed from the diet.
(2008). Previous studies have demonstrated a clear effect Increased consumption of plant material as a result of
of plant Si on a range of insect feeding guilds, includ- poor dietary quality has been reported for Sp. eridania
ing lepidopteran borers (e.g. Chilo suppressalis Walker (Cramer) (Lepidoptera: Noctuidae) (Peterson et al., 1988)
(Crambidae) (Nakano et al., 1961), Ch. infuscatelus Snell and Schis. gregaria (Massey et al., 2006). However, silica
(Crambidae) (Rao, 1967), Diatraea saccharalis (Fabri- may also alter the palatability of plant material and deter
cius) (Crambidae) (Anderson & Sosa, 2001), Sesamia herbivore feeding, as shown by Massey et al., (2006) for
calamistis Hampson (Noctuidae) (Sétamou et al., 1993), Schis. gregaria and Sp. exempta fed on grass species in which
Cn. medinalis (Sudhakar et al., 1991), Scirpophaga excerptalis uptake of Si produced increased leaf abrasiveness.
Walker (Pyralidae) (Gupta et al., 1992) and E. saccharina Other studies have also shown reduced consumption
(Kvedaras et al., 2007a,b; Kvedaras & Keeping, 2007), when feeding on high-Si plants. Salim & Saxena (1992)
folivores (e.g. Spodoptera exempta Walker (Lepidoptera: observed decreased rice consumption by white-backed
Noctuidae) and Schistocerca gregaria Forskål (Orthoptera: planthopper, Sogatella furcifera (Horvath) (Homoptera:
Acridoidea) (Massey et al., 2006), phloem-feeding insects Delphacidae) fed on susceptible rice cultivars high in Si,
(e.g. Schiz. graminum (Rondani) (Hemiptera: Aphididae) in addition to reduced growth, adult longevity, fecundity
(Basagli et al., 2003; Moraes et al., 2004), Laodelphax and population increase. Chu & Horng (1991) noted that

174 Ann Appl Biol 155 (2009) 171–186 © 2009 The Authors
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O.L. Reynolds et al. Silicon and insect herbivores

slag-treated corn plants had harder stems, and consump- were applied in artificial diet, rather than in planta,
tion of the leaves (from Si-treated and untreated plants) thereby removing the natural context under which
by Asiatic corn borer Ostrinia furnacalis was negatively the insect would encounter Si when feeding on the
correlated with tissue hardness, indicating that Si-based plant. Similarly, Zouhourian-Saghiri et al.’s (1983) study,
leaf hardness may be a factor in resistance to this borer. although quantitative, did not employ Si treatments (and
However, stem and sheath consumption was not affected untreated controls) of the same plant material to exclude
by the treatments. A reduction in stalk damage and lar- the likely contribution of cellulose and lignin (within
val weight gain (as a result of decreased consumption) their high-Si plant material, Sasa japonica) to mandibular
was shown for stem borer E. saccharina fed on Si-treated wear in Locusta migratoria (Orthoptera: Acrididae).
sugarcane compared with untreated sugarcane (Keeping Redmond & Potter (2007), showed that calcium silicate
& Meyer, 2006; Kvedaras et al., 2007b). fertiliser (Excellerator™, Excell Minerals, Sarver, PA)
In Ost. furnacalis Guenée (Lepidoptera: Crambidae) applied to ’Penncross’ creeping bentgrass caused no
reared on artificial diet, mortality of larvae and pupae excessive wearing of mandibular teeth in black cutworm,
increased, pre-oviposition period was prolonged and the Agrotis ipsilon (Hufnagel) (Lepidoptera: Noctuidae) and
oviposition period was shortened as the diet Si content root-feeding masked chafer grubs, Cyclocephala spp.
was increased from 0.1, to 3, 5 or 10% (Horng & Chu, (Coleoptera: Scarabaeidae). There are only two studies
1990). Pupal weight, fecundity, net reproductive rate and that have attempted to accurately quantify the effect of
intrinsic rate of increase were all negatively correlated experimentally elevated plant Si on mandibular wear
with diet Si content. However, the duration of larval and in insects. Kvedaras et al., (2009b) showed only a non-
pupal stages, longevity of adults and mean generation significant trend for greater mandibular wear in E.
times were not significantly affected.
saccharina larvae fed on sugarcane cultivars treated with
Silicon may counteract the effects of high plant N levels
Si compared with untreated controls. However, Massey
in promoting insect performance. In potted corn (Zea mays
& Hartley (2009) demonstrated a significant and rapid
L.), Chu & Horng (1991) observed decreased preference
(within a single instar) increase in mandibular wear for
under free-choice conditions of Ost. furnacalis for slag-
Sp. exempta fed on two out of three grass species fertilised
treated (i.e. Si-treated) plants but increased preference for
with Si, compared with untreated controls. A difficulty in
high-nitrogen-treated plants. Meyer & Keeping (2005)
measuring mandibular wear in Lepidoptera is that larvae
later showed that the application of soil-applied Si
replace their mandibles at each moult. Plants high in Si
can mitigate the promotional effects of applied N on
may force larvae to moult sooner than usual, because
populations of the stalk borer, E. saccharina in sugarcane.
of increased mandibular wear, which may in turn lead
They postulated that the use of Si would enable growers,
to decreased body weight. This is an area that merits
who had reduced their N application rates because of
additional study, as the time spent in a given instar
increased E. saccharina infestations, to resume applying
would probably affect the degree of mandibular wear and
the recommended rates of N, thus ensuring that N would
not limit the crop yield. resulting long-term performance of the herbivore.
Recently, Hunt et al., (2008) concluded that silica may
defend grasses, at least in part, by mechanical protection
Physical defences of resources in the chlorenchyma cells (which contain
In larval Lepidoptera, mandibular wear is often attributed high levels of starch and protein sought after by insects),
to plant Si (sometimes based only on visual comparisons through reduced mechanical breakdown of the leaf. They
between high-Si and low-Si plants or tissue) and has been demonstrated that high-silica grasses released less chloro-
reported for rice stem borer Ch. suppressalis (Sasamoto, phyll after grinding and retained more after passing
1958; Djamin & Pathak, 1967; Dravé & Laugé, 1978) through the gut of the locust, Schis. gregaria, illustrating
and leaf folder larvae Cn. medinalis (Hanifa et al., 1974; that Si levels are correlated with increased mechanical
Ramachandran & Khan, 1991) fed on rice and for fall protection.
armyworm, Sp. frugiperda, fed on corn plants (Goussain Using scanning electron microscopy and X-ray micro-
et al., 2002). However, these studies and most other analysis Keeping et al., (2009) showed that in two sug-
reports of Si causing mandibular wear of arthropods have arcane cultivars, Si-treated plants had increased silica in
shortcomings (Smith et al., 2007; Kvedaras et al., 2009b) the stalk epidermis, particularly at the internode and root
and are therefore not compelling in their conclusions. band. As these are known penetration sites for E. sac-
For example, although Dravé & Laugé (1978) used charina borer, such patterns of Si deposition may partly
a quantitative, statistical approach to compare larval explain the enhanced resistance of Si-treated sugarcane
mandibular wear of Ch. suppressalis, their Si treatments to borer penetration and feeding. However, Keeping et al.

Ann Appl Biol 155 (2009) 171–186 © 2009 The Authors 175
Journal compilation © 2009 Association of Applied Biologists
Silicon and insect herbivores O.L. Reynolds et al.

(2009) conceded that fibre content (cellulose, hemicel- Furthermore, the authors showed that the elevated Si
lulose and lignin) is probably also crucial in this regard concentration in grass leaves as a result of herbivory
(more so in resistant cultivars; Rutherford et al., 1993). deterred further feeding. These results are not surprising
In maize, for example, Coors (1987) demonstrated that as it is recognised that artificial damage can elicit different
genotypes most resistant to European corn borer Ost. induction responses to those of herbivory, and in insects,
nubilalis (Hübner) had high levels of total structural car- salivary gland secretions and regurgitants have been
bohydrates, lignin and silica. identified as the sources of cues leading to plant responses
Although numerous studies have looked at the direct to herbivory (see review by Felton & Eichenseer, 1999).
physical effects of Si on insect herbivores (see above),
there have been few studies that have looked at the Location and composition of Si in the plant
indirect effects. Kvedaras & Keeping (2007) showed that
Si delayed the penetration of the sugarcane stalk by E. Several studies suggest that the site and arrangement of
saccharina, which they proposed may lead to increased Si deposition in the plant are more important than Si
exposure time of young larvae to adverse environmental content per se in restricting larval feeding. Miller et al.,
conditions or control practices that target such larvae. (1960) demonstrated that the leaf sheaths of certain
Similarly, when yellow stem borer, Sci. incertulas, larvae wheat cultivars, as well as one cultivar of oats, which
were fed on rice grown in a high Si nutrient solution, are resistant to hessian fly Phytophaga destructor (Say)
penetration time increased compared with a low level of (Diptera: Cecidomyiidae), displayed a more complete
Si (IRRI, 1990, cited by Savant et al., 1997). Feeding and even distribution of silica deposits on their surface
and boring by Ch. suppressalis larvae is also impeded than those of susceptible cultivars (Miller et al., 1960).
in rice grown in silica (Djamin & Pathak, 1967). A The authors proposed that this arrangement may allow
further indirect effect (and to our knowledge, the only P. destructor larvae to feed between the rows of Si and
example to date of an interaction between Si-mediated showed that in certain resistant cultivars there may not
resistance to an insect herbivore and an abiotic stress be sufficient free space to allow unobstructed feeding
factor) was illustrated by a reduction in E. saccharina by larvae. Blum (1968) found that in five cultivars of
numbers and stalk damage in Si-fed but water-stressed sorghum resistant to the sorghum shoot fly, Atherigona
cane plants (Kvedaras et al., 2007a). Further, the increase soccata (Rondani) (Diptera: Muscidae), the density of
in resistance of Si-treated water-stressed susceptible silica bodies present in the abaxial epidermis at the
cultivars to E. saccharina was such that borer survival base of the first, second and third leaf sheaths (the
and damage in these plants approached, and in almost area where the insect penetrates the plant) was greater
all instances, was not significantly different from that of than in a susceptible cultivar. Similarly, a study on rice
resistant cultivars (irrespective of whether the latter were revealed that the pattern of silica in the epidermis of
treated with Si and/or water stressed). Possibly, in the cultivars resistant to the leafroller, Cn. medinalis comprised
presence of Si, water-stressed borer-susceptible cultivars closer silica chains, heavier deposition of silica in the
may develop a defensive chemistry with a profile similar intercostal area, higher epidermal silica deposition, and
to that of borer-resistant cultivars. These results parallel single or double rows of silica in comparison with
other studies, where the effect of Si on plant disease is susceptible cultivars (Hanifa et al., 1974). As the total
more pronounced under conditions of abiotic stress than Si content of resistant and susceptible cultivars did not
non-stressed conditions (Ma et al., 2001; Ma, 2004; Gong differ significantly, the physical arrangement and location
et al., 2005; Wiese et al., 2005). Such observations suggest of Si in the plant tissues were considered to be important
that Si application could provide enhanced herbivore (Miller et al., 1960; Hanifa et al., 1974). Barker (1989)
resistance to plants exposed to a range of abiotic factors, showed that high densities of silica deposits (inclusive of
including salinity stress and heavy metal toxicity. trichomes) in the sheath epidermis of ryegrass cultivars
In both an insect (the locust, Schis. gregaria) and a hindered oviposition by the weevil, Listronotus bonariensis
mammalian herbivore (the field vole, Microtus agrestis (Kuschel) (Coleoptera: Curculionidae). The closely spaced
L. (Rodentia: Muridae), Massey et al., (2007) revealed rows of silica cells in the leaf epidermal layer is thought
that repeated damage events led to the induction of to be partly responsible for the greater resistance of
increases of c. 400% in foliar Si levels of two grass species, wild rice to Cn. medinalis compared with the wider
compared with undamaged plants and attributed this to spacing in a rice hybrid (Ramachandran & Khan, 1991).
either increased Si uptake from the soil or increased A rice cultivar resistant to white-backed plant hopper,
deposition of phytoliths, or both. Single damage events S. furcifera not only had higher concentrations of Si but
or mechanical defoliation, however, were not sufficient silicated bulliform cells were closer and double in number
to promote induction responses in either grass species. compared with a highly susceptible cultivar (Mishra &

176 Ann Appl Biol 155 (2009) 171–186 © 2009 The Authors
Journal compilation © 2009 Association of Applied Biologists
O.L. Reynolds et al. Silicon and insect herbivores

Misra, 1992). In a study comparing ryegrass cultivars resulting in a reduction of probing time, leading them
(hybrid type of Zoysia japonica Steud. with Z. matrella to conclude that chemical changes because of Si absorp-
(L.) Merr., cv. Miyako) resistant and susceptible to larvae tion by the plant were likely responsible, as opposed to
of Osc. frit, Moore (1984) observed that the resistant a physical impediment, as the stylet eventually did reach
cultivar had silica bodies scattered over the pseudostem, the phloem. Studies on Si-mediated resistance of cucum-
but in the susceptible cultivar they were often in ber to whitefly B. tabaci (Correa et al., 2005) and wheat
discrete rows. This may explain why small increases to Schiz. graminum (Gomes et al., 2005) also suggest that
in plant silica levels correspond to large reductions in soluble Si is important in induced resistance to insect
the number of larvae found (Moore, 1984). Kvedaras herbivores, as has been shown for Si-mediated resis-
et al., (2007b) suggested this could also be true for tance induction of various crops to fungal pathogens (see
E. saccharina on sugarcane. Rao & Panwar (2001), in review by Fauteux et al., 2005). Research on the inter-
comparing maize cultivars, either resistant, moderately actions between plant defences, Si and fungal pathogens
resistant or highly susceptible to shoot fly species A. soccata has shown that the presence of a pathogen is pivotal
and A. naqvii Steyskal, showed that the lignified vascular in the mobilisation of biochemical defences and in the
bundles and leaf epidermal silica bodies were negatively upregulation of defence-related genes; some of the lat-
correlated with oviposition and dead-heart percentages, ter are involved in the biosynthesis of herbivore-induced
and that the resistant cultivars had a higher number of plant volatiles (HIPVs), including jasmonate (JA) and
lignified vascular bundles as well as leaf epidermal silica salicylate (SA) (Thaler et al., 2002; Fauteux et al., 2005,
bodies compared to susceptible cultivars. Agarwal (1969) 2006). HIPVs are released in response to herbivore dam-
reported that sugarcane Saccaharum spontaneum L. clones age to facilitate location by natural enemies of plants
with higher numbers of silica cells in the wax band of where their prey or hosts are present (see review by
the internode had significantly lower levels of sugarcane Dicke et al., 2003 and later section on tri-trophic inter-
scale, Melanaspis glomerata (Green) (Homoptera: Coccidae) actions). Work in the USA has shown that plant-derived
infestation. or synthetic versions of these chemical cues (HIPVs)
will attract beneficial insects into treated crops (Khan
et al., 2008). Silicon, either alone or together with Schiz.
Chemical defence
graminum pre-infestation, elicited a significant increase
As early as 1958, Sasamoto suggested that host choice by in the defensive enzymes, peroxidase, polyphenoloxi-
an insect depended not only on the physical properties dase, and phenylalanine ammonia-lyase (PAL), activity
of the food but also on its chemical properties. In a in wheat (Gomes et al., 2005). Peroxidase is involved in
laboratory choice study, Sasamoto (1958) demonstrated the process of lignification and suberin synthesis, which
that larvae of the rice stem borer, Ch. suppressalis displayed increase the hardness of tissues and in the production
a preference for untreated rice stalks over Si-treated of quinones and active oxygen that possess antibiotic
rice stalks, and that larvae from the latter showed properties (Goodman et al., 1986; Bowles, 1990; Stout
increased mandibular wear (see previous section). et al., 1994). Polyphenoloxidase catalyses the oxidation of
Notably, water extracted from the stem of the rice plant phenolic compounds to quinones that leads to a reduc-
cultured in nitrogen-rich manure was more attractive tion in the nutritional quality of the food and decreased
to larvae than the silicated one. Recently, Zadda et al., protein digestibility (Felton & Duffey, 1990; Felton et al.,
(2007) showed that the application of organic sources 1994). The PAL enzyme increases the production of phe-
of nutrients and amendments (farm yard manure, nolic compounds that have anti-nutritional, deterrent
poultry manure, neem cake, mahua cake, pungam and toxic properties (Appel, 1993). Many mechanisms
cake and biofertilisers, i.e. Azospirillum, phosphobacteria of defence are shared with those against fungi (Goussain
and silica solubilising bacteria) to eggplant, Solanum et al., 2005), and as the bulk of the work thus far on
melongena L., significantly enhanced the production Si and induced chemical defence has been carried out
of defensive chemicals including silica and phenols. in this area, it may be that future research will reveal
Further, the authors showed that induced resistance similar defensive mechanisms at work in both fungi and
by way of antibiosis led to a reduction in feeding insects.
rate, oviposition, longevity and population buildup and
prolonged the nymphal duration of eggplant pests
Nil effects of Si on herbivores
(Zadda et al., 2007).
Earlier, Goussain et al., (2005) showed that stylet pene- In India, Agarwal (1969), using morphological studies,
tration of Schiz. graminum was not impeded by Si in wheat was unable to demonstrate any relationship between
plants; however, the stylet was withdrawn more often the number of silica cells in sugarcane clones and

Ann Appl Biol 155 (2009) 171–186 © 2009 The Authors 177
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Silicon and insect herbivores O.L. Reynolds et al.

infestation by whitefly Aleurolobus barodensis Mask. at present there is no convincing evidence (Keeping &
and Neomaskellia bergii Sign. (Homoptera: Aleyrodidae). Kvedaras, 2008).
Application of calcium silicate to five different turf grasses
did not negatively affect the growth and development
Silicon, solubilisers and other control agents
of tropical sod webworm, Herpetogramma phaeopteralis
Guenée (Lepidoptera: Pyralidae), despite increased Si It has been demonstrated that molecules such as pyridine-
levels in the plant tissue (Korndörfer et al., 2004). Prilled N-oxides can enhance the solubility of Si in water and the
calcium silicate fertiliser (Excellerator™, Excell Minerals, subsequent physiological effects on yellow stem borer,
Sarvar, PA) applied to fairway-height turf on silt loam Sci, incertulas (Walker) (Lepidoptera: Pyralidae) damage
soil elevated leaf Si by as much as 40% without reducing and rice blast disease, Magnaporthe grisea (Ranganathan
the palatability or suitability for the black cutworm, A. et al., 2006). However, more recently Voleti et al., (2008)
ipsilon, nor did Si reduce the density or weight of root- showed, using scanning electron microscopy and Si
feeding masked chafer (Cyclocephala spp.) grubs (Redmond mapping, that simple amino acids (natural Si solubilisers)
& Potter, 2007). Similarly, sodium silicate drenches that such as histidine or glycine can significantly enhance the
elevated leaf Si content of greenhouse-grown bentgrass levels of Si(OH)4 in the stem and increase Si deposition in
from 0.5 to 2.5% did not reduce A. ipsilon feeding or the leaf. This is important progress because pyridine-N-
survival and had little effect on larval growth rates oxides are organic molecules that may have soil residual
(Redmond & Potter, 2007). effects harmful to soil microorganisms, unlike the natural
Chaudhary & Yadav (1998) found no correlation Si solubilisers identified above. Subsequently, a novel
between the incidence of top borer, Sc. excerptalis and class of biocompatible molecules has been identified
the presence of silica, cellulose, lignin and ash present in that exhibit remarkable resistance to damage caused by
midribs, growing points and leaf blades of 30 sugarcane the yellow stem borer, Sci. incertulas and blast infections
genotypes, despite a large variation in all but the lignin in rice, in addition to high levels of dry matter and
contents of midribs, growing points and leaf blades. increased yields. In a free-choice test, Si, the insect growth
In International Rice Research Institute and new plant regulator lufenuron, and their interaction did not affect
type (NPT) rice cultivars, Sunio et al., (2000) could not the preference of Sp. frugiperda on maize; however, when
show a correlation between silica content and percentage both were combined in a non-choice test, better control
whiteheads and percentage dead hearts caused by stem of Sp. frugiperda was obtained compared with lufenuron
borers (mainly Ch. suppressalis). Although Hanisch (1981) only (Neri et al., 2005). Tomquelski et al., (2007) showed
showed that infestation of nitrogen-fertilised wheat plants that both acibenzolar-S-methyl (BTH) and Si applied
by Si. avenae and M. dirhodum was reduced, through independently to cotton, reduced the larval period and
application of a 1% sodium silicate foliar spray, to pupal weight and increased the pupal period of Alabama
below that in nitrogen-deficient control plants, apparently argillacea (Hübner) (Lepidoptera: Noctuidae). Recently,
because of the uptake of silica in the leaves of sprayed Gomes et al., (2008a) demonstrated that the combined
plants, Massey et al., (2007) found that the same phloem- use of Si and a half dose of the insecticide imidacloprid
feeding insect, Si. avenae, suffered no detrimental effects reduced the colonisation of potato plants by the aphid,
of feeding on five different grasses with naturally high Myzus persicae (Sulzer) to the same extent as the insecticide
levels of Si. alone. This prompted the authors to propose that Si
Mebrahtu et al., (1988) could not demonstrate any application could potentially form part of an Integrated
correlation between the silicon content of soya beans and Pest Management (IPM) program for potatoes. A further
the pupal weight of the Mexican bean beetle, Epilachna study that year showed that although Si fertilisation did
varivestis Mulsant (Coleoptera: Coccinellidae). In three not affect M. persicae preference, it did reduce the fecundity
pineapple varieties, a significant positive correlation was and the rate of population growth of the insects (Gomes
found between Si and the population density of the et al., 2008b).
pineapple flat mite, Dolichotetranychus floridanus (Banks)
(Acarina: Tenuipalpidae) (although not an insect, but
Silicon and plant cultivars
a member of the Phylum Arthropoda) (Das et al.,
2000). There may be many more studies that remain Although the bulk of our discussion has focussed on Si
unpublished on the failure of Si to reduce insect herbivore amendments, the discovery of the association between
performance, as these are often regarded as not worthy endogenous levels of plant Si and resistance to insects has
of publication, but would assist us in identifying any led the way for applied Si amendments to enhance plant
trend in how plant Si may differentially affect insects resistance to insect herbivores. Elevated plant Si levels
belonging to different groups or feeding guilds, for which or higher densities of Si bodies are frequently reported

178 Ann Appl Biol 155 (2009) 171–186 © 2009 The Authors
Journal compilation © 2009 Association of Applied Biologists
O.L. Reynolds et al. Silicon and insect herbivores

from plant cultivars that have high or increased levels of BTH, which reduced oviposition more than three times
resistance to insect attack. This has been demonstrated that of the control. Both application methods of calcium
for aubergine (brinjal/eggplant) (Jat & Pareek, 2003), silicate and BTH resulted in increased developmental
barley (Guslits, 1990; Sinel’nikov & Shapiro, 1990), period of second to fourth instar nymphs. Interestingly,
cotton (Taneja et al., 1988), cotton and okra (Singh & Costa & Moraes (2006) showed that the application of Si
Agarwal, 1988), chickpea (Rupali et al., 2003), maize (foliar- and soil-applied, although particularly the latter)
(Sekhon & Kanta, 1997; Smith, 1997; Rao & Panwar, or BTH, significantly reduced the number of nymphs, the
2001), citrus (orange) (Matichenkov et al., 2000), rice population growth rate, the post-reproductive period and
(Sujatha et al., 1987; Anuradha et al., 1988 a,b; Dan & the longevity of Schiz. graminum on wheat plants, leading
Chen, 1990; Mishra et al., 1990; Sudhakar et al., 1991; the authors to conclude that either would be encouraging
Mishra & Misra, 1992, 1993; Lin, 1993; Soliman et al., control methods in the IPM of the aphid in wheat. In
1997; Saeb et al., 2002), sorghum (Narwal, 1973; Chavan a later study on the same species, Costa et al., (2007)
et al., 1990; Dalvi et al., 1990), sugarcane (Kennedy & compared foliar Si application, BTH and both combined,
Nachiappan, 1992; Keeping & Meyer, 2006), sweet potato and found that the latter two were less preferred by
(Singh et al., 1993) and turmeric (Lakshmi & Sudhakar, aphids than the foliar Si-treatment alone.
2003). Clearly, however, Si is only one factor that acts Some studies, whether using soil and/or foliar-applied
to enhance cultivar resistance to insects, along with a Si, have failed to measure plant Si levels following the
multitude of other physical and chemical traits that have treatments to establish the extent of Si uptake (Basagli
been documented in the plant resistance literature. et al., 2003; Moraes et al., 2004; Gomes et al., 2005;
Goussain et al., 2005). Such measurements are essential
in supporting causal arguments that the applied Si is
Foliar- versus root-applied Si
acting on the insect through incorporation into the plant
A number of studies have showed increased plant tissue, as opposed to some extraneous pathway, and are
resistance to biotic stress after the application of Si foliar especially important with respect to foliar applications.
sprays. Hanisch (1981) reported that the deposition and Matlou (2006) concluded that foliar application of three
increased solubility of Si in the leaves of wheat plants after sources of Si (including potassium silicate) at two rates
foliar sprays of a 1% Na2 SiO2 solution were responsible and including the use of a wetting agent was ineffective
for the resistance of this crop to the aphids M. dirhodum in increasing leaf Si content of sorghum (a Si-responsive
and Si. avenae. The author showed that the number of hair crop) grown in low-Si soils, whereas the root-applied
cells containing silica in the epidermis of plants treated calcium silicate treatment produced significant increases
with sodium silicate was 70% greater than in untreated in leaf Si. Guével et al., (2007) showed that at least for
ones, indicating the uptake and translocation of silica wheat plants Si is absorbed primarily, if not exclusively,
in the leaves of foliar sprayed plants. Foliar application by the root system. They noted that foliar sprays with
of lignite fly ash dust, which is high in silica content, both Si and nutrient solutions are likely to have a direct
to papaya led to a significant reduction in the disease, effect on the plant, rather than one mediated by the plant
papaya leaf curl virus, and in vector, silverleaf whitefly, as is the case for root amendments. However, this area
B. tabaci populations (Eswaran & Manivannan, 2007). remains contentious as there is no firm evidence for the
Studies have also compared the application of Si uptake of Si from the leaves (Guével et al., 2007).
as both a foliar spray and via the soil. For example,
Moraes et al., (2005) showed that the number of corn
Tri-trophic interactions
leaf aphids, Rhopalosiphum maidis (Fitch) (Hemiptera:
Aphididae) on detached leaves of corn leaf plants was Plants supplemented with Si translocate silicic acid
significantly reduced when the plants had been treated throughout their tissues and, when attacked by
with either two foliar Si sprays or one application of pathogens, produce systemic stress signals such as SA
Si through the soil and an additional Si foliar spray and JA (Fauteux et al., 2005) that are key to plant-
compared with plants treated with Si applied through the induced defences (Gatehouse, 2002). Jasmonic acid, a
soil or one foliar Si spray. naturally occurring growth regulator and wound hor-
In a free-choice study on whitefly B. tabaci (Hemiptera: mone in higher plants, is particularly important because
Aleyrodidae), Correa et al., (2005) reported that calcium of its role in HIPV production. Silicon may have an effect
silicate incorporated in the soil resulted in a reduced on the plant’s ability to generate an induced response by
number of whitefly eggs on cucumber; however, the acting not only at the second but also the third trophic
most striking effect was that of a foliar application of level (i.e. by attracting predators or parasitoids). However,
calcium silicate with or without the resistance inducer only one study has tested for the effect of Si on an insect

Ann Appl Biol 155 (2009) 171–186 © 2009 The Authors 179
Journal compilation © 2009 Association of Applied Biologists
Silicon and insect herbivores O.L. Reynolds et al.

herbivore and the resultant attractancy of the insect’s difficult to demonstrate and there is still considerable
predators and parasitoids (Moraes et al., 2004). That work scope in this area for worthwhile research.
showed no effect of Si on natural enemies (detrimen- Even less well explored is the likely role of soluble Si
tal effects on the pest were shown however), but used in induced plant defence against insect herbivores. The
non-choice conditions in which parasitoid wasps, Aphid- limited research that has been done in this area provides a
ius colemani Viereck (Hymenoptera: Aphidiidae) were tantalising glimpse of how Si may be involved in induced
restricted on a small scale to individual wheat plants biochemical defences that call into play various defensive
that were narrowly spaced. Further experiments con- enzymes and defence-related plant hormones such as
ducted with the predator, Chrysoperla externa (Hagen) JA and SA. It has become clear that Si is critical as an
(Neuroptera: Chrysopidae) were still less favourable for ameliorator of plant stress, whether biotic or abiotic, with
the detection of induced plant defences involving HIPVs, possibly little role in unstressed plants. The importance
as the herbivore Schiz. graminum was removed from the of (drought) stress in amplifying the protective effect
test plants and fed to predators that had not been exposed of Si against a stem borer has been demonstrated, and
to plants at all (Moraes et al., 2004). this, together with clues provided by studies of ind-
Work recently commenced in Australia, presented at uced biochemical defence in Si × pathogen interactions
the IV Silicon in Agriculture Conference by Kvedaras (Menzies et al., 1992; Chérif et al., 1994; Fauteux et al.,
et al., (2008), and since published (Kvedaras et al., 2005) and two studies on Si and resistance induction
(2009a) has been designed to look at the attraction of against insects in wheat and cucumber (Gomes et al.,
natural enemies to Si-treated and untreated plants and 2005; Kvedaras et al., 2009a), poses the questions: (a) Is
is methodologically and conceptually more advanced. soluble Si a key player in facilitating induced biochemical
The application of Si with a subsequent pest infestation defences against insect herbivores? (b) Is Si involved in
increased the plants’ ability to mount an induced response upregulation of defence-related genes (such as those in
by attracting natural enemies. An effect that was reflected JA or SA biosynthesis) following insect attack? (c) While
in elevated biological control in the field. The potential insect attack is a source of stress for the plant, do
for using Si fertilisation as an elicitor of induced plant additional abiotic stresses in combination with Si lead
chemical defences for attraction of biological control to an even greater inducible response?
agents in pest management is an exciting prospect. These are just a few of the many questions that will
Moreover, if the promising results to date translate to no doubt emerge as the field of Si-mediated defence of
the availability of novel plant protection compounds that plants against herbivores (invertebrate and vertebrate)
promote host plant resistance traits operating through progresses. Throughout this review we have attempted to
the third trophic level it may prove to be less prone to a highlight some of the novel hypotheses and findings thus
decrease in efficacy as a result of genetic adaptation of the far, while also indicating the shortcomings of research
target pest population (Gurr & Kvedaras, 2009). This is so in the area of Si and insect herbivory. We hope that
as pest suppression is likely to occur through a number this review may assist in providing pointers for further
of natural enemies capable of adapting in response to any investigation into the challenging new areas of this field
shift in pest phenotype. that remain little explored or unexplored.

Acknowledgements
Conclusions and future prospects
Lee-Anne McInerney is thanked for a literature search
Although the discovery that Si plays a role in the resis- and both Christine Harley and Lee-Anne McInerney
tance of plants to insect herbivores dates back to the early are thanked for sourcing many of the more obscure
1950s, we are only in the very early stages of elucidating references. Fumi Chiku is thanked for the interpretation
the mechanisms on which this is based. It seems fairly of the Japanese language articles and Ana Korndörfer
clear that Si-mediated defences against insects are partly for interpretation of Brazilian articles. We are grateful
physical in nature and therefore based on solid amor- to Geoff Gurr who provided useful comments on
phous (opaline) Si deposited in key tissues and organs earlier drafts of the manuscript and to two anonymous
of the plant, where they are able to mechanically hinder reviewers.
insect establishment, plant penetration and feeding effi-
ciency, as well as to reduce palatability and plant tissue
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