Sci. Agri. 8 (2), 2014: 80-85 © PSCI Publications Scientia Agriculturae www.pscipub.com/SA E-ISSN: 2310-953X / P-ISSN: 2311-0228 DOI: 10.15192/PSCP.SA.2014.4.2.8085 The Ameliorative effects of silicon element on improvement of plants tolerance to diseases Vahid Ghazi Mohseni1 and Seyed Kazem Sabbagh2 1. MSc student of plant pathology, University of Zabol, Iran 2. Assistant professor, faculty of plant pathology, University of Zabol, Iran Corresponding author email: [email protected] Paper Information Received: 16 September, 2014 Accepted: 20 October, 2014 Published: 20 November, 2014 ABSTRACT Silicon is the second most plentiful element in soil and is beneficial for a large variety of plants. It is concentrated in plant tissues in quantities similar to that of macronutrients. Considerable damages to plants caused by abiotic stresses such as drought stress, salinity stress, heavy metal stress and nutrient imbalance, as well as biotic stresses like insect pests and pathogens and even herbivorous attacks, have been reported to be reduced significantly by silicon application. It seems that better understanding about the function of silicon in plant would be help to improve production in areas with soils poor in silicon, since it has been known to have the ability to lessen vulnerability of plant against different kinds of diseases. Si provides many benefits to stressed plants including protection against attack from certain fungal, bacterial and virus pathogens. The studies have shown that silicon fertilization to be efficacious in controlling and mitigating severity of some diseases such as rice blast, powdery mildew, bacterial wilt, bacterial speck Tobacco ringspot virus and tobacco mosaic virus. Thus, more knowledge about silicon function can help farmers and agriculture industry to manage and control diseases effectively. © 2014 PSCI Publisher All rights reserved. Key words: disease, plant, silicon, yield Introduction The silicon (Si) is an abundant element in terrestrial superficie (Pereira et al., 2003), however its availability to plants is normally low (Hattori et al., 2005). According to Matichenkov and Calvert (Matichenkov and Calvert, 2002), the chemically active Si forms in soil are represented by soluble monosilicic acid (Si (OH)4) that is soluble and weakly adsorbed, as well as acid polisilicic, which are compound organosilicates. Si is considered an benefic element to higher plants (Epstein and Bloom, 2004), being that the absorption process must be active or passive (Ma et al., 2007), and deposition in cell walls of several organs such as leaf and stem can promote beneficial effects (Cunha et al., 2008), and for this reason has been frequently linked to physiological, morphological, nutritional, and molecular aspects in plants (Isa et al., 2010; Ma et al., 2004; Ma and Yamaji, 2006; Lobato et al., 2009). In plants this nutrient is assimilated mainly by roots, and capacity to accumulate in tissues is variable (Chiba et al., 2009), being several monocots such as Oryza sativa and Triticum aestivum considered silicon accumulator, with absorption active by root system, and it present leaf levels normally higher that 10.0 g kg-1 of Si (Oliveira, 2009). In other hand, many dicots like as Phaseolus vulgaris and Glycine max are characterized as not accumulator of silicon, and its presents passive absorption, with leaf tenors minors that 5.0 g kg-1 of Si (Takahashi et al., 1990). In tissues, about of 99% of silicon is found in silic form and less than 1% is colloidal or ionic form, which is the soluble form (Ma et al., 2001). Therefore, the storage sites of silicon in plants normally are responsible to improve leaf and plant architectures and also others metabolic processes like as gas exchanges (Pereira et al., 2012), photosynthetic pigments (Silva et al., 2012), and antioxidant system (Li et al., 2012), in which it results in better performance linked to growth, development, and yield parameters (Nolla et al., 2012). The positive impacts of silicon in improving plant growth and yield are significant, and these include increasing resistance against environmental stresses such as cold and salinity stress, insect attacks and penetration of pathogens (Jones and Handreck, 1967; Elawad and Green, 1979; Belanger et al., 1995; Savant et al., 1996). In addition to what has been mentioned above, using appropriate amount of silicon has been found to reduce the probability of plants being grazed by herbivores (Rodrigues and Datnoff, 2005; Belanger et al., 1995; Datnoff et al., 1997; Savant et al., 1996). Meanwhile, a large number of valuable investigations have been conducted on the positive effects of silicon in plants (Abed Ashtiani et al., 2012). Sci. Agri. 8 (2), 2014: 80-85 Typically in soils with low amounts of silicon, silicon fertilization has been applied to increase both the quality and quantity of agricultural crops such as rice and sugar cane (Korndorfer and Lepsch, 2001). In addition, foliar applications of silicon have been confirmed to increase the resistance against pathogens in plant species that do not absorb silicon efficiently (Epstein, 1994; Bowen et al., 1992). Recent research has demonstrated that both foliar and soilborne diseases of cucumber and other cucurbits can be suppressed by applying this element (Belanger et al., 1995). Much of the research on silicon has been focused on rice, although its benefits to other crops such as wheat and corn have also been proven (Wang et al., 2001). The goal of this paper is to review the research findings in relation to silicon and plant diseases. Silicon deficiency in the soil Silicon (Si) is one of the most abundant elements in the earth’s crust and most soils contain considerable quantities of the element (Epstein, 1994). However, repeated cropping can reduce the levels of plant available Si to the point that supplemental Si fertilization is required for maximum production and some soils contain little plant-available Si in their native state. Low-Si soils are typically highly weathered, leached, acidic and low in base saturation. Thus, highly weathered soils such as Oxisols and Ultisols can be quite low in soluble Si (Foy, 1992). Highly organic Histosols that contain little mineral matter may contain little Si. Interestingly, soils comprised mainly of quartz sand (SiOz) such as sandy Entisols also may be very low in plant-available Si. These conditions are found in many crop-producing areas of Africa, Asia, Latin America and the southeastern USA. Genotype and silicon interactions Effective disease-control strategies for agronomic crops, in particular, include the incorporation of genetically controlled resistance into cultivars. Since genotypes vary in disease resistance, the relationship between Si content among genotypes and disease resistance was investigated. In a test of 18 rice cultivars grown at three locations representing high (116 mg Si L-1 soil), typical (40 mg Si L-1 soil) and low (6 mg Si L-1 soil) soil Si status, cultivars varied significantly for tissue Si concentration (Deren et al., 1992). Certain genotypes consistently ranked high or low in Si concentration across all locations. This suggested that the acquisition of Si may be an inherited trait. Similar variations in Si concentration have been reported for African japonicas (Winslow, 1992). Genotypes were further evaluated for Si accumulation and brown spot development on a low-Si soil fertilized with 0 and 2 Mg Si ha-1 (Deren et al., 1994). Genotypes varied significantly for Si concentration in plant tissue. However, the range among genotypes was very narrow. Rico 1 and Della X2 tended to have the greatest Si concentrations in both Si-deficient and Si-amended soils (Datnoff et al., 1997). The suppressive effect of silicon on fungal diseases Rice blast (Pyricularia oryzae) Several researchers have demonstrated that silicon application has a suppressive effect on fungal diseases such as rice blast (Magnaporthe grisea, teleomorph, Pyricularia oryzae, anamorph), brown spot (Cochliobolus miyabeanus, teleomorph; Bipolaris oryzae, anamorph), leaf scald (Gerlachia oryzae), sheath blight (Rhizoctonia solani, teleomorph) and stem rot (Sclerotium oryzae, teleornorph) (Datnoff et al., 1991, 1997; Savant et al., 1997; Seebold, 1998; Winslow, 1992). Under silicon deficient conditions, some diseases such as blast, brown spot and sheath blight can be extremely threatening to rice cultivation (Rodrigues and Datnoff, 2005). Accumulated silicon in rice tissues enhances resistance against insects and diseases, increases erectness of leaves resulting in increased photosynthesis, improves water usage, and decreases toxicity of heavy metals and cuticular transpiration (Nakata et al., 2008; Epstein, 1994). The amounts of silicon accumulated in rice plants can be as much as 10% of plant dry weight. It is often several times higher than the rate of accumulation of other essential macro nutrients such as nitrogen, phosphorus and potassium (Nakata et al., 2008; Ma et al., 2006). Rice blast disease, caused by the fungus Pyricularia oryzae (Couch and Kohn, 2002), is particularly prevalent in all areas under rice cultivation and it also causes heavy yield losses (Nakata et al., 2008; Ou, 1985). Development of efficient and cost-effective methods to manage the disease is one of the priority issues in rice production (Hayasaka et al., 2008). There are many management methods that are being evaluated for rice blast disease management and these include, among others, breeding of resistant varieties (time consuming) and management methods to improve soil nutrients (Hayasaka et al., 2008). The minimum amount of silicon needed to control blast disease in rice is 3-5% (Datnoff et al., 1997). Positive effects and importance of silicon as a nutrient element in rice plants have been reported in several studies (Hayasaka et al., 2008; Ando et al., 2002; Ma et al., 2006; Kamenidou et al., 2009; Deren et al., 1992; Ma and Takahashi, 2002; Gao et al., 2004) and a gene which plays a role in the transport of silicon has also been detected (Hayasaka et al., 2008; Ma et al., 2006). There are different hypotheses on how silicon confers and induces resistance to some plants specially rice against diseases such as blast (Ishiguro, 2001). These hypotheses can be classified into two groups (Hayasaka et al., 2008). One hypothesis is based on the belief that silicon acts as a mechanical barrier against fungal penetration (Hayasaka et al., 2008) which is also known as physical resistance, while the second hypothesis is based upon physiological resistance. 81 Sci. Agri. 8 (2), 2014: 80-85 Powdery mildew (Golovinomyces cichoracearum) Gerbera (Gerbera jamesonii) is among the major greenhouse grown cut flowers in the United States. Powdery mildew caused by Golovinomyces cichoracearum (syn. Erysiphe cichoracearum) is one of the most destructive diseases of gerbera (Malathrakis and Goumas, 1999). Golovinomyces cichoracearum forms white powdery blotches on leaves and other green parts of gerbera, resulting in increased water loss, reduced photosynthesis and, consequently, reduced quality of flowers. In commercial situations, control of powdery mildew on gerbera has involved frequent applications of fungicides. Environmentally friendly, effective alternatives to fungicides for management of powdery mildew are deemed desirable. Due to the Food Quality Protection Act (FQPA) of 1996, many fungicides currently used to control diseases in floriculture crops are at risk of de-registration by the Environmental Protection Agency (EPA). In addition, powdery mildew fungi can develop resistance to conventional fungicides. Therefore, research is needed to identify alternatives to conventional fungicides. A promising alternative to conventional fungicides for control of powdery mildew on gerbera is the element silicon (Si). In highly relevant work conducted in Florida, applications of Si (calcium silicate) resulted in dramatic reductions of two foliar fungal diseases of rice (Datnoff et al., 1991). Although calcium silicate contains other nutrients, only Si increased within plant tissues and correlated with disease reduction. In some cases, Si use provided disease control equivalent to that of fungicides in both rice and turf (Datnoff et al., 2001 and Brecht et al. 2004). In addition, fungicide rates could be reduced when used in combination with this element (Seebold et al., 2004). More recently, Seebold et al (2000) and Rodrigues et al (2001) demonstrated that host resistance of susceptible and partially resistant cultivars can be augmented with Si to the same general level as those containing complete disease resistance. When a Si source such as calcium silicate is incorporated in potting media low in plant available Si, Si is taken up by plants via the root system at higher rates than other elements and enhances plant growth as well as the defense response of plants to pathogen attack. Recent research demonstrated that Si can play a role in enhancing the growth and appearance of flowering ornamentals. Savvas et al. (2002) showed that Si in the nutrient solution resulted in significantly thicker flower stems and a higher proportion of flowers graded Class I for gerbera (Gerbera jamesonii). Shoot number, flower number, dry weight were enhanced by 25, 62 and 66 %, respectively, over the control for paper daisies (Helichrysum adenohorum) amended with Si (Muir et al., 1999). In a similar study, open flowers per head were increased by 28% in snapdragons while bent/lodged stems were reduced by 33%. More recently, black spot infection of Rosa hybrida ‘Meipelta’ was demonstrated to be suppressed by the addition of 150 mg/L Si as potassium silicate to irrigation water (Gillman et al., 2003). The role of silicon on plant responses to the bacterial diseases Bacterial wilt (Ralstonia solanacearum) Diseases caused by different fungal and bacterial pathogens (Jones et al., 1991) are among the major constraints of tomato production. Bacterial wilt caused by Ralstonia solanacearum (Yabuuchi et al., 1995) is one of the most important diseases that limit tomato production worldwide. The disease also constitutes a serious damage to the cultivation of many other Solanaceous crops (potato, tobacco, pepper and eggplant) in tropical, sub-tropical and temperate regions (Hayward, 1991). Different management strategies of bacterial wilt disease have been studied in many parts of the world with special focus on host plant resistance, cultural practices, and soil amendments (Saddler, 2005). Silicon (Si) has been reported for having a significant effect in reducing bacterial wilt incidence in tomato in a hydroponics culture system, and peat substrate (Dannon and Wydra, 2004). Diogo and Wydra (2007) also reported an induced basal resistance on cell wall level, in silicon treated tomato plants compared to non-treated plants. Thus, an elucidation of the effect of silicon amendment under field conditions, for the management of bacterial wilt on tomato is necessitated. Ayana et al. (2011) have reported that silicon fertilizer significantly reduced the bacterial population, mean wilt incidence, percent severity index, and corresponding areas of disease incidence, and severity progress curves in the moderately resistant tomato cultivar. However, silicon fertilizer do not resulted in any significant reduction of all disease parameters in the moderately susceptible cultivar Marglobe. Bacterial speck (Pseudomonas syringae) Bacterial speck caused by Pseudomonas syringae pv. tomato (Pst) is an economically important disease on tomato crops (Silva and Lopes, 1995). Yield losses of up to 30% are reported in environments with high relative humidity and temperature ranging from 20 to 25ºC (Silva and Lopes, 1995). The control of bacterial diseases is limited to the availability of chemicals. Products containing copper and the inducer of resistance acibenzolar-S-methyl are the most used to control bacterial speck (Lopes and Santos, 1994). Also, the use of cultivars containing the resistant Pto gene, such as the cultivar BRS Tospodoro, is highly recommended to the growers (Lopes and Santos, 1994). Other control methods for bacterial speck remain to be investigated. Silicon (Si) has been gaining attention in the control of certain fungal diseases, such as powdery mildew in muskmelon and tomato (Dallagnol et al., 2012; Yanar et al., 2011). Dannon and Wydra (2004) obtained reduction in the 82 Sci. Agri. 8 (2), 2014: 80-85 symptoms of bacterial wilt (Ralstonia solanacearum) in tomato plants from susceptible and moderately resistant genotypes supplied with Si. According to Diogo and Wydra (2007), the reduction in the symptoms of bacterial wilt occurred due to changes in cell wall structure of the xylem vessels in plants supplied with Si. In the same pathosystem, Ghareeb et al. (2011) found that the expression of genes related to basal defense of tomato was induced in plants supplied with Si and inoculated with R. solanacearum, thereby, demonstrating the importance of Si in enhancing host defense responses. Effect of silicon on virus diseases Tobacco ringspot virus and tobacco mosaic virus Si provides many benefits to stressed plants including protection against attack from certain fungal and bacterial pathogens. For example, treatment of tomato varieties and Cucumis sativus with Si reduced infection by Ralstonia solanacearum and Podosphaera xanthii, respectively (Dannon and Wydra, 2004). While Si protects plants against fungal and bacterial pathogens, the effects of this element on viral infections are unclear. In an experiment in order to investigate the role of Si in viral infections, hydroponic studies were conducted in Nicotiana tabacum with two pathogens: Tobacco ringspot virus (TRSV) and Tobacco mosaic virus (TMV). Plants grown in elevated Si showed a delay in TRSV systemic symptom formation and a reduction in symptomatic leaf area, compared to the non-supplemented controls. TRSV-infected plants showed significantly higher levels of foliar Si compared to mock-inoculated plants. However, the Si effect appeared to be virusspecific, since the element did not alter TMV symptoms nor did infection by this virus alter foliar Si levels. Hence, increased foliar Si levels appear to correlate with Si-modulated protection against viral infection. This is all the more intriguing since N. tabacum is classified as a low Si accumulator (Zellner et al., 2011). Conclusion Although the function of silicon in plant is not wholly recognized, the application of this element would be an encouraging method to improve production in areas with soils poor in silicon, since it has been known to have the ability to lessen vulnerability of plant against different kinds of diseases, specifically those caused by fungal pathogens such as Pyricularia oryzae. An important fact about silicon in addition of the aforementioned points is that it is an environmentally friendly element in relation to soils, fertilizers and plant nutrition. Hence, it can be beneficial in IPM programs for those crops where silicon has been proven to have a positive impact. The positive effects of silicon in plant nutrition and resistance against diseases have been well established. Thus, providing the knowledge about silicon to farmers and rice growers will help the agriculture industry to manage and control rice diseases effectively. 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