Contents Summary 67 I. Introduction 68 II. Silicon transport in plants: to absorb or not to absorb 69 III. The role of silicon in plants: not just a matter of semantics 71 IV. Silicon and biotic stress: beyond mechanical barriers and defense priming 76 V. Silicon and abiotic stress: a proliferation of proposed mechanisms 78 VI. The apoplastic obstruction hypothesis: a working model 79 VII. Perspectives and conclusions 80 Acknowledgements 81 References 81 Summary: Silicon (Si) is not classified as an essential plant nutrient, and yet numerous reports have shown its beneficial effects in a variety of species and environmental circumstances. This has created much confusion in the scientific community with respect to its biological roles. Here, we link molecular and phenotypic data to better classify Si transport, and critically summarize the current state of understanding of the roles of Si in higher plants. We argue that much of the empirical evidence, in particular that derived from recent functional genomics, is at odds with many of the mechanistic assertions surrounding Si's role. In essence, these data do not support reports that Si affects a wide range of molecular‐genetic, biochemical and physiological processes. A major reinterpretation of Si's role is therefore needed, which is critical to guide future studies and inform agricultural practice. We propose a working model, which we term the ‘apoplastic obstruction hypothesis’, which attempts to unify the various observations on Si's beneficial influences on plant growth and yield. This model argues for a fundamental role of Si as an extracellular prophylactic agent against biotic and abiotic stresses (as opposed to an active cellular agent), with important cascading effects on plant form and function.

Silicon (Si) is a bioactive element associated with beneficial effects on mechanical and physiological properties of plants. Silicon alleviates abiotic and biotic stresses, and increases the resistance of plants to pathogenic fungi. Several studies have suggested that Si activates plant defense mechanisms, yet the exact nature of the interaction between the element and biochemical pathways leading to resistance remains unclear. Silicon possesses unique biochemical properties that may explain its bioactivity as a regulator of plant defense mechanisms. It can act as a modulator influencing the timing and extent of plant defense responses in a manner reminiscent of the role of secondary messengers in induced systemic resistance; it can also bind to hydroxyl groups of proteins strategically involved in signal transduction; or it can interfere with cationic co-factors of enzymes influencing pathogenesis-related events. Silicon may therefore interact with several key components of plant stress signaling systems leading to induced resistance.

▪ Abstract The controlled environment of greenhouses, the high value of the crops, and the limited number of registered fungicides offer a unique niche for the biological control of plant diseases. During the past ten years, over 80 biocontrol products have been marketed worldwide. A large percentage of these have been developed for greenhouse crops. Products to control soilborne pathogens such as Sclerotinia, Pythium, Rhizoctonia and Fusarium include Coniothyrium minitans, species of Gliocladium, Trichoderma, Streptomyces, and Bacillus, and nonpathogenic Fusarium. Products containing Trichoderma, Ampelomyces quisqualis, Bacillus, and Ulocladium are being developed to control the primary foliar diseases, Botrytis and powdery mildew. The development of Pseudomonas for the control of Pythium diseases in hydroponics and Pseudozyma flocculosa for the control of powdery mildew by two Canadian research programs is presented. In the future, biological control of diseases in greenhouses could predominate over chemical pesticides, in the same way that biological control of greenhouse insects predominates in the United Kingdom. The limitations in formulation, registration, and commercialization are discussed, along with suggested future research priorities.

The role and essentiality of silicon (Si) in plant biology have been debated for >150 years despite numerous reports describing its beneficial properties. To obtain unique insights regarding the effect of Si on plants, we performed a complete transcriptome analysis of both control and powdery mildew-stressedArabidopsisplants, with or without Si application, using a 44K microarray. Surprisingly, the expression of all but two genes was unaffected by Si in control plants, a result contradicting reports of a possible direct effect of Si as a fertilizer. In contrast, inoculation of plants, treated or not with Si, altered the expression of a set of nearly 4,000 genes. After functional categorization, many of the up-regulated genes were defense-related, whereas a large proportion of down-regulated genes were involved in primary metabolism. Regulated defense genes included R genes, stress-related transcription factors, genes involved in signal transduction, the biosynthesis of stress hormones (SA, JA, ethylene), and the metabolism of reactive oxygen species. In inoculated plants treated with Si, the magnitude of down-regulation was attenuated by >25%, an indication of stress alleviation. Our results demonstrate that Si treatment had no effect on the metabolism of unstressed plants, suggesting a nonessential role for the element but that it has beneficial properties attributable to modulation of a more efficient response to pathogen stress.

Silicon (Si) amendments in the form of exogenously supplied nutrient solution or calcium silicate slag protect wheat plants from powdery mildew disease caused by the fungus Blumeria graminis f. sp. tritici. The most striking difference between Si- and Si+ plants challenged with B. graminis f. sp. tritici was the extent of epidermal cell infection and colonization by B. graminis f. sp. tritici. Histological and ultrastructural analyses revealed that epidermal cells of Si+ plants reacted to B. graminis f. sp. tritici attack with specific defense reactions including papilla formation, production of callose, and release of electron-dense osmiophilic material identified by cytochemical labeling as glycosilated phenolics. Phenolic material not only accumulated along the cell wall but also was associated with altered integrity of haustoria in a manner similar to localized phytoalexins as reported from other pathosystems. These results strongly suggest that Si mediates active localized cell defenses against B. graminis f. sp. tritici attack.