Silicon (Si) is one of the most abundant elements in the earth's crust, although its essentiality in plant growth is not clearly established. However, the importance of Si as an element that is particularly beneficial for plants under a range of abiotic and biotic stresses is now beyond doubt. This paper reviews progress in exploring the benefits at two‐ and three‐trophic levels and the underlying mechanism of Si in enhancing the resistance of host plants to herbivorous insects. Numerous studies have shown an enhanced resistance of plants to insect herbivores including folivores, borers, and phloem and xylem feeders. Silicon may act directly on insect herbivores leading to a reduction in insect performance and plant damage. Various indirect effects may also be caused, for example, by delaying herbivore establishment and thus an increased chance of exposure to natural enemies, adverse weather events or control measures that target exposed insects. A further indirect effect of Si may 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.

1 The stalk borer Eldana saccharina is the most destructive pest in sugarcane in South Africa. This study investigated: (1) the potential of applied silicon in enhancing plant resistance of sugarcane to E. saccharina, using calcium silicate as a carrier; (2) whether there was any interaction between cane variety (and stalk borer resistance) and silicon treatment. 2 Six commercial varieties of sugarcane were treated in a pot-plant trial with two levels (5000 and 10 000 kg/ha) of calcium silicate. After artificial infestation with E. saccharina, response to the treatments was assessed in terms of borer numbers and mass, and stalk damage. 3 Calcium silicate significantly enhanced resistance at the higher rate compared with the control. Borer mass was reduced by 19.8% and stalk length bored by 24.4%. Lower treatment values were intermediate between those of the higher treatment and the control. 4 The interaction between variety and Si treatment was not significant when varieties were examined individually in the analysis. However, the interaction was significant for borer mass when varieties were grouped according to their resistance characteristics. Susceptible varieties might benefit more from treatment with silicon than resistant ones, as resistant varieties showed no significant effect of silicon. 5 All varieties had increased silicon content due to the treatments, but differed appreciably in stalk silicon content at the different treatment levels. Similarly, within varieties, stalk silicon content did not correspond consistently with borer response patterns and silicate application rates.

Many plants grown in soils amended with silicon (Si) display increased levels of resistance to attack by insect herbivores. This study aimed to determine if Si treatment impeded Eldana saccharina Walker (Lepidoptera: Pyralidae) stalk penetration and subsequent damage, as well as borer mass gain, on the node and internode of a susceptible (N11) and a resistant (N33) sugarcane cultivar. Sugarcane [Saccharum spp. (Poaceae)] cultivars were grown in a pot trial in Si‐deficient river sand, with (Si+) and without (Si–) calcium silicate. Sugarcane was infested with 2‐week‐old E. saccharina larvae and harvested at four times, 24, 48, 72, and 96 h after infestation. Silicon‐treated plants showed significant increases in Si content compared to controls, and the external rind was significantly harder for Si+ cane than Si– cane. Silicon treatment significantly decreased borer penetration, stalk damage, and larval mass gain. The results are consistent with the hypothesis that Si contributes to sugarcane stalk borer resistance by impeding larval penetration. Silicon appears to contribute to the suppression of E. saccharina directly through reduced larval growth and feeding damage to the crop, and indirectly by delaying stalk penetration, resulting most likely in increased exposure time of young larvae to natural enemies, adverse climatic factors, or control measures that target young larvae (e.g., insecticides).

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1 Silicon can increase the resistance of plants to attack by herbivorous insects. The present study aimed to determine the effect of silicon and cultivar on mandibular wear in larvae of the sugarcane stalk borer Eldana saccharina Walker (Lepidoptera: Pyralidae). 2 Four sugarcane cultivars, resistant (N21, N33) and susceptible (N11, N26) to E. saccharina were grown in a pot trial in silicon deficient river sand, with (Si + ) and without (Si -) calcium silicate. Individual third-instar larvae were confined on the sugarcane stalk at three known feeding sites (leaf bud, root band and internode) and left to feed for 21 days. 3 Eldana saccharina larval heads were mounted on stubs, with the mandibles oriented horizontally and photographed under a scanning electron microscope. Mandibular wear was measured from the digital images using a quantitative method. 4 Although there was a trend for increased wear in larvae that developed on Si + cane, no significant effect of silicon, cultivar or site on mandibular wear of E. saccharina was shown. 5 This is the first study to accurately and quantitatively measure the mandibular wear of an insect fed on Si + plants.

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Silicon (Si) can improve resistance of plants to insect attack and may also enhance tolerance of water stress. This study tested if Si-mediated host plant resistance to insect attack was augmented by water stress. Four sugarcane cultivars, two resistant (N21, N33) and two susceptible (N26, N11) to Eldana saccharina Walker were grown in a pot trial in Si-deficient river sand, with (Si+) and without (Si-) calcium silicate. To induce water stress, irrigation to half the trial was reduced after 8.5 months. The trial was artificially infested with E. saccharina eggs after water reduction and harvested 66 days later. Silicon treated, stressed and non-stressed plants of the same cultivar did not differ appreciably in Si content. Decreases in numbers of borers recovered and stalk damage were not associated with comparable increases in rind hardness in Si+ cane, particularly in water-stressed susceptible cultivars. Overall, Si+ plants displayed increased resistance to E. saccharina attack compared with Si- plants. Borer recoveries were significantly lower in stressed Si+ cane compared with either stressed Si- or non-stressed Si- and Si+ cane. Generally, fewer borers were recovered from resistant cultivars than susceptible cultivars. Stalk damage was significantly lower in Si+ cane than in Si- cane, for N21, N11 and N26. Stalk damage was significantly less in Si+ combined susceptible cultivars than in Si- combined susceptible cultivars under non-stressed and especially stressed conditions. In general, the reduction in borer numbers and stalk damage in Si+ plants was greater for water-stressed cane than non-stressed cane, particularly for susceptible sugarcane cultivars. The hypothesis that Si affords greater protection against E. saccharina borer attack in water-stressed sugarcane than in non-stressed cane and that this benefit is greatly enhanced in susceptible cultivars is supported. A possible active role for soluble Si in defence against E. saccharina is proposed.