Abstract
Silicon is the second most abundant element and accounts for 27–28% of the earth’s crust. Biological systems also contain significant amounts of silicon, as amorphous silica (SiO2·nH2O), and its soluble form, silicic acid (Si(OH)4). Plant dry biomass contains 0.1–10% of silicon. Despite its extensive distribution in plants, silicon is viewed as a quasi-essential element, as most of the plant species can live their entire life in its absence. Interestingly, even in higher amounts, silicon is harmless, noncorrosive, and nonpolluting to plants. It is typically accumulated in the epidermal cells, creating external dual layers of silica-cuticle and silica-cellulose on leaves, stem, and hulls. It thus acts as a physical barrier in plants. Si also alleviates the stress-induced responses in plants. This chapter reviews how plants benefit from silicon under adverse environmental conditions.
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1 Introduction
Food security is the most important fundamental need of society. The wide-ranging increase in environmental damage and the pressure of ever-increasing human population have adversely affected global food production (Etesami and Jeong 2018). The world population today is estimated to be about 7 billion and projected to reach between 7.5 to 10.5 billion by 2050 (Godfray et al. 2010). Such an enormous rise in the population would demand higher agricultural productivity per unit area from already degraded lands. Moreover, climate change has aggravated the occurrence and intensities of various biotic and abiotic stresses (Etesami and Jeong 2018). Such conditions would compel farmers to cultivate marginal lands and poor soils (Glick 2014).
Stress affects the growth and development of the plants, thereby leading to more significant losses in agricultural productivity. However, plants have adopted numerous mechanisms to tolerate stress and survive stress-induced conditions. Healthy plants are capable of combating stress, and plant nutrients are vital to maintaining healthy plant growth. The microelements or micronutrients are known to give stress tolerance to plants (Vanderschuren et al. 2013; Bradáčová et al. 2016). Though the roles of several macro- and micronutrients in plants have been well documented, few of the nutrient elements have remained neglected. This chapter focuses on the role of silicon, one of the neglected plant nutrients, and its role in plants suffering from adverse environmental conditions.
2 Adverse Environmental Conditions
2.1 Biotic Stress
Throughout their life, plants get exposed to a multitude of stresses that modify plant growth and development. Organisms like fungi, bacteria, mycoplasma, insets, nematodes, weeds, and parasitic plants induce biotic stress. The viruses and viroids, though nonliving, also contribute to the biotic stress. These agents affect the plant growth and development by depriving nutrients leading to reduced plant vigor and death of plants in extreme cases (Das and Rakshit 2016). The severity of biotic stress depends on the environmental factors, cropping systems, types of crops, cultivars, and resistance levels of plants. Hot and humid conditions and poor crop management practices are the two leading causes of biotic stresses (Pantazi et al. 2020). Early recognition of biotic stress is the key to control it via integrated pest management and the use of pesticides.
Plants do not have an adaptive immune system like vertebrates. They can neither adapt to new diseases nor memorize the previous infections. However, plants have developed several mechanisms to combat biotic stresses. They rely on various physical and chemical barriers that confer strength and rigidity to survive under biological stress.
2.2 Abiotic Stress
The nonliving factors imposing adverse effects on healthy growth and development of the plants are called abiotic stresses. These include drought, salinity, heavy metals, too low or too high temperatures, and other environmental extremes. These factors can reduce the crop yield by 51–82% (Bray et al. 2000). Plants combat these stresses at various levels like morphological, physiological, biochemical, and molecular levels (Husen 2010; Getnet et al. 2015; Embiale et al. 2016; Husen et al. 2016, 2018, 2019; Hussein et al. 2017; Siddiqi and Husen 2017, 2019; Zeng et al. 2020; Kar and Öztürk 2020). Over the past few decades, advances in plant physiology, genetics, and molecular biology have greatly upgraded our understanding in terms of crops respond to stress conditions. These responses depend not only on their duration and severity but on the age and the developmental stage of the plant as well (He et al. 2018).
3 Is Silicon Essential to Plants?
Silicon (Si) is the eighth-most abundant element in the universe. In earth’s crust, its abundance ranks only second to oxygen. The lithosphere contains about 27.7% silicon (Epstein 1999). It rarely occurs in its pure form, and more than 90% of the Si in the earth’s crust exists as silicates (Mitra 2015).
Biological systems also contain significant amounts of silicon, as amorphous silica (SiO2·nH2O), and its soluble form, silicic acid (Si(OH)4). The first indication of in vivo formation of organosilicates, their distribution, and physiological importance was discovered in a diatom Navicula pelliculosa (Kinrade et al. 2002). Plants also contain significant amounts of Si that can range from 0.1 to 10% on the dry weight basis (Epstein 1994; Ma and Takahashi 2002; Hodson et al. 2005; Ma et al. 2006). Differences in the levels of silicon in different plants could be due to the differential ability of roots to absorb Si (Takahashi et al. 1990). Despite its high amounts in plants, Si is looked upon as a quasi-essential element since most of the plant species can live their entire life in the absence of silicon (Arnon and Stout 1939). Nonavailability of Si-free environment due to its contamination in purified water, chemicals, and dust might be the reasons for considering Si as nonessential for higher plants (Liang et al. 2015). Therefore, adhering to the definition of essentiality proposed by Epstein and Bloom (2005), Si is a quasi-essential element in plants.
Interestingly, there are several reports on the positive roles of Si in the plant growth (Eneji et al. 2008; Soundararajan et al. 2014; Zhang et al. 2015), yield (Epstein 1999), structural toughness (Epstein 1994), nutrient management (Tripathi et al. 2012), and absorption of light (Li et al. 2004). Its role in accelerating the tolerance to biotic and abiotic stresses in plants has also been explained (Ma 2004; Cookson et al. 2007; Liang et al. 2007; Muneer et al. 2014; Soundararajan et al. 2014). How Si alleviates biotic and abiotic stresses has become a booming topic of interest. In the past 15 years, several researchers have reported and reviewed the positive effects of Si under biotic and abiotic stresses (Fig. 1a, b). However, studies on Si in conjunction with abiotic stress were significantly more than those with biotic stress (Fig. 1b). This chapter summarizes how plants use silicon and respond to Si availability during adverse environmental conditions.
(a) Silicon-related publications in the plant sciences from 2005 to 2020 (Till June) (Based on PubMed search with the keywords “silicon” and “abiotic stress”). (b) Silicon-related publications in the plant sciences from 2005 to 2020 (Till June) (based on PubMed search with the keywords “silicon” and “biotic stress”)
4 Uptake of Si in Plants Under Adverse Environmental Conditions
In plants, roots take up more than 90% of Si and translocate it to shoots (Ma and Takahashi 2002). Roots absorb Si in the form of silicic acid at pH < 9 (Takahashi and Hino 1978; Raven 2001; Ma and Takahashi 2002). The concentration of silicic acid in soil solutions usually ranges between 0.1 and 0.6 mM (Epstein 1994), and in some cases, up to 0.8 mM (Sommers and Lindsay 1979). The soil pH modulates the solubility of silicates, and with the increasing pH, solubility decreases. However, most of the crops are cultivated in soils with pH well below the alkaline mark of pH 9.0.
Studies in rice have shown that roots take up Si from the rhizosphere by some kind of transporter, which transports it radially from the root cortical cells to the xylem (Tamai and Ma 2003). Once absorbed, Si is transported to the shoot as silicic acid. In different plants like rice, cucumber, and tomato, the concentration of silicic acid in the root cell symplast was higher than that in the external solution. Rice has shown a significantly higher concentration of Si than observed in cucumber and tomato (Mitani et al. 2005). This difference in the ability to take up Si is attributed to the different modes of transport in these plants. Moreover, the xylem loading of Si in rice occurs through a transporter. In contrast, in cucumber and tomato, it occurs by passive diffusion.
Plants differ in their ability to take up and distribute Si. The highest levels of Si uptake are reported in bryophytes and lycopods and Equisetum among the pteridophytes. However, ferns and gymnosperms tend to accumulate Si in lesser quantities (Takahashi et al. 1990). Two of the angiosperm families, viz., Cyperaceae and Poaceae, are known to accumulate Si at higher concentrations (Hodson et al. 2005). Depending upon their ability to accumulate Si, plants are categorized into three classes: Si accumulators (e.g., rice, wheat, millet, and sugarcane) since they absorb large quantities of Si; Si non-accumulators (e.g., Snapdragon); and Si excluders (e.g., soybean) (Van der Vorm 1980; Marschner 1995).
So far, only a few genes have been identified that are involved in the uptake of Si in plants. The first of those genes is Lsi1 that was reported in rice (Ma et al. 2006). The Lsi1 gene is expressed mainly in roots, and its encoded protein has Si transporter activity. The Lsi1-encoded protein is located on the distal sides on plasma membranes in both the layers of exodermis and endodermis. Bioinformatics tools have revealed that the Lsi1 belongs to a subfamily of aquaporin Nod26-like major intrinsic proteins (NIP). Chiba et al. (2009) have reported the HvLsi1 gene in barley for the influx of Si from roots. The presence of ZmLsi1 and ZmLsi6 transporter from maize was reported by Mitani et al. (2009). Bokor et al. (2017) have studied the expression of ZmLsi1, ZmLsi2, and ZmLsi6 genes and their effects on Si uptake and ionome content in maize (Bokor et al. 2017).
5 Transport of Si in Plants
Si absorbed by the root cells must be transported to other plant organs. Therefore, Si must be taken out of the root cells first. In marine organisms, the influx and efflux of Si are mediated by the same protein (Hildebrand et al. 1998). In rice, however, the efflux is mediated by a transporter Lsi2 (Ma et al. 2006), and Lsi6 mediates the influx of silicic acid from xylem to xylem parenchyma cells, thus influencing the distribution of Si in rice roots (Yamaji et al. 2008).
Plants are capable of synthesizing Si-rich molecules of various sizes. The accumulated Si provides rigidity and roughness to the plant cell walls (Epstein and Bloom 2005) and also offers other beneficial effects (Van Soest 2006). The passive transport of Si driven by transpiration also leads to its deposition on the cell wall. Researchers have used several biophysical tools like scanning electron microscopy (SEM) coupled with X-ray microanalysis, laser ablation (LA), X-Ray fluorescence spectrometry, and X-ray absorption near edge structure (XANES) to study the distribution of Si in plants (Rufo et al. 2014; Bokor et al. 2017). Si is mostly deposited in the epidermal cells of leaves, stems, and hulls where double layers of silica-cuticle or silica-cellulose containing hydrated polymers of amorphous silica are formed on their surface (Fauteux et al. 2005; Wiese et al. 2007; Deshmukh et al. 2017).
6 Role of Si in Plants Under Adverse Environmental Conditions
It is speculated that the global climate changes will trigger more frequent incidences of biotic and abiotic stresses leading to severe agricultural losses. The abiotic stresses reduce the global agricultural yield by as much as 70% (Acquaah 2012). How to fulfill the ever-increasing food demand under such circumstances will be a real challenge. The application of Si in soils deteriorated due to abiotic stresses has been beneficial for crop productivity. A summary of the beneficial and or positive effects of Si in plants exposed to various biotic and abiotic stresses is presented in Table 1.
6.1 Si and Plant Growth
Seed germination plays a significant role during seedling establishment. Drought adversely affects seed germination leading to agriculture losses (Hubbard et al. 2012; Shi et al. 2014). However, there are few reports on the effects of Si on seed germination under drought stress (Hameed et al. 2014; Shi et al. 2014). Priming of wheat seeds with sodium silicate was beneficial in enhancing the rate of seed germination under drought stress (Hameed et al. 2014). Similar observations were reported in tomato (Siddiqui and Al-Whaibi 2014), and maize (Zargar and Agnihotri 2013) seeds germinated under drought stress.
All the essential nutrients are required in adequate amounts for the healthy growth and development of plants. The process of absorption of these nutrients from the surrounding is disturbed under various stresses (Gunes et al. 2007a; Chen et al. 2011; Khattab et al. 2014). The deposition of Si in the endodermal layer of root cells helps in the selective uptake of nutrients, and such deposition reduces the accumulation of toxic ions in different plant parts (Yeo et al. 1999). The soil application of Si has enhanced the uptake of macronutrients (P, K, Ca, and Mg) and micronutrients (Fe, Cu, and Mn) in sunflower (Gunes et al. 2008a). The application of Si to the rice plants subjected to drought stress showed an increase in the uptake of potassium and phosphorus (Khattab et al. 2014). An increased levels of phosphorus (Gong and Chen 2012), and potassium and calcium (Kaya et al. 2006) were observed in wheat under drought stress. In other grasses such as Chloris gayana, Sorghum sudanense, Festuca arundinacea, and Phleum pratense, the levels of N, P, and K were increased upon the application of Si under drought stress (Eneji et al. 2008).
6.2 Effect of Si on Structure and Physiology of Plants
Si plays two critical roles under adverse environmental conditions: physical and mechanical protection due to its deposition in the epidermal layer, and triggering a biochemical response to metabolic changes. Numerous researchers have reported the deposition of Si in the form of phytoliths in plant tissues (Katz 2015). Evidence of cross-linking of Si in cell walls with hemicellulose is also reported (He et al. 2015; Luyckx et al. 2017). Si accumulates in the epidermal layer of leaves in the form of silica bodies. This deposition of Si in various forms improves mechanical properties and may act as a physical barrier (Massey et al. 2007). Such Si deposition might also increase roughness and tensile strength of leaves, causing reduced palatability and digestibility in herbivores (Massey and Hartley 2009; Hartley et al. 2015; Frew et al. 2016).
The supplementation of Si has proven beneficial to reduce the transpirational loss of water from leaves (Gong et al. 2003). It also enhanced the UV tolerance that resulted in reduced membrane damage (Goto et al. 2003; Shen et al. 2010). Stomatal conductance in relation to turgidity in guard cells is also reduced due to the deposition of Si in leaves (Zhu and Gong 2014). Under drought stress, plants can absorb water from the soil due to Si-induced root elongation and upregulation of aquaporin genes in roots (Hattori et al. 2005; Liu et al. 2015). The supply of Si reduces the translocation of toxic ions such as Na+, Cl−, and heavy metals from root to shoot (Savvas and Ntatsi 2015). Si-containing materials alter the rhizospheric pH and limit the bioavailability of heavy metals (Wu et al. 2013). In contrast, soluble silicates produce metasilicic acid (H2SiO3), which is gelatinous and retains heavy metals (Gu et al. 2011).
6.3 Role of Si in Plant Defense Under Adverse Environmental Conditions
Supplementation of Si fertilizers enhances the defense mechanisms of plants against pathogens such as viruses, bacteria, fungi, and other organisms like nematodes, arthropods, vertebrates, and herbivores (Griffin et al. 2015; Reynolds et al. 2016). Si mitigates the biotic stress in plants by either acting as a physical barrier in the epidermal layer or by alleviating resistance to pathogens. The distribution of silica in the leaf tissues can contribute more to the defense against herbivorous insects than other animals (O’Reagain and Mentis 1989). Likewise, the deposition of phytoliths throughout the leaf epidermis acts as a barrier against leaf-chewing insects than the phloem-feeding insects (Massey et al. 2006).
The application of Si improves the plant’s ability to restrict the spread of pathogens. For example, enhanced resistance to Eldana saccharina in sugarcane was examined by Keeping et al. (2009). Si application has also reduced the rates of infections by pathogenic fungi such as Rhizoctonia solani and Bipolaris oryzae (Ning et al. 2014; Schurt et al. 2014; Zhang et al. 2014). In the Si-supplemented wheat plants, the invasion by Pyricularia oryzae and Bipolaris sorokiniana was restricted within the leaf epidermis (Domiciano et al. 2013).
6.4 Effect of Si on the Plant Biochemical Responses Under Adverse Environments
At the biochemical level , Si contributes to the defense mechanisms by increasing the levels of diverse secondary metabolites like phenolics, flavonoids, momilactones, and phytoalexins (Cherif et al. 1994; Rémus-Borel et al. 2005; Debona et al. 2017). It also enhances the activities of defense enzymes like chitinase, lipoxygenase, peroxidase, phenylalanine ammonia-lyase, and polyphenol oxidase (Rahman et al. 2014). Signaling of key phytohormones like salicylic acid, jasmonic acid, and ethylene that are active during stress is also influenced by the Si treatments (Glazebrook 2005; Wu and Baldwin 2010; Liang et al. 2015). Si also interferes with the insect’s life cycle by lowering the phenology, thereby making it more prone to predation (James 2003; Connick 2011). Elevated malondialdehyde (MDA) contents reflect membrane damage caused due to lipid peroxidation (Zhu et al. 2004). The MDA levels were reduced upon supplementation of Si in barley (Liang et al. 2003), grapevine (Soylemezoglu et al. 2009), and maize (Moussa 2006). Additionally, Si also influences the levels of osmolytes and plant growth regulators (Adrees et al. 2015; Ali et al. 2015; Noman et al. 2015; Jabeen et al. 2016).
7 Si and Osmolytes
Osmolytes are organic solutes that maintain the cellular potential for a healthy metabolism. They do not interfere with the normal metabolism of the plants (Zhang et al. 2004) but protect the cellular enzymes and cell membranes from the detrimental effects of high ion concentrations due to stress (Bohnert and Shen 1999; Ashraf and Foolad 2007). Thus, osmolytes act as osmoprotectants and include low molecular weight solutes like glycine betaine (GB), proline, polyols, alanine betaine, and simple sugars like trehalose and sucrose (Sharma et al. 2019). These solutes help the host to sustain severe osmotic stress (Singh et al. 2015) by maintaining the osmotic balance between the cytosol and surrounding medium of the cell. Osmolytes are also known to inhibit the production of ROS, thereby protecting the plants from oxidative damage. Plants produce osmolytes mainly under adverse environmental conditions, especially abiotic stress. Accumulation of osmolytes indicates the plant’s adaptation to stress.
Si seems to modulate the levels of osmolytes in stressed plants. Application of Si reduced the proline levels in stressed plants of spinach and tomato (Gunes et al. 2007b), wheat (Tuna et al. 2008), sorghum (Yin et al. 2013), soybean (Lee et al. 2010), and grapevine (Soylemezoglu et al. 2009). The levels of glycine betaine, proline, and total soluble sugars were elevated after foliar application of Si in the tolerant as well as sensitive okra genotypes exposed to salt stress. However, the effect was more pronounced in sensitive genotypes (Abbas et al. 2015). A similar trend was also reported in capsicum (Pereira 2013), tobacco (Pereira 2013), and maize (Sayed and Gadallah 2014). Exposure of Si has enhanced the plant tolerance to drought stress via osmolytes modification in many crops (Crusciol et al. 2009), such as the augmented proline content in drought-stressed condition for wheat (Gong et al. 2005; Kaya et al. 2006) and pepper plants (Pereira 2013).
8 Si and Phytohormones
Phytohormones induce the vital responses needed for the healthy growth and development of plants. Apart from their regulatory functions, they also coordinate signal transduction pathways under biotic and abiotic stress (Wolters and Jürgens 2009). Si application regulates the levels of phytohormones to enhance plant tolerance to stress (Kim et al. 2014). However, the level of ethylene declined after the application of Si under salinity stress in sorghum (Yin et al. 2016). In soybean, the level of GA was elevated, and that of abscisic acid (ABA) declined in the presence of Si (Lee et al. 2010). Similarly, the level of jasmonates (JA) was reduced, and that of salicylic acid (SA) increased in the presence of Si (Hamayun et al. 2010). Si induced the thermo-tolerance in potato by regulating the endogenous level of SA and ABA. The Si-mediated tolerance to brown spot disease in rice depends on immune hormones SA and JA as well as fungal ethylene (Van Bockhaven et al. 2015). Likewise, Si priming of seeds gave tolerance to powdery mildew in A. thaliana (Vivancos et al. 2015).
9 Si and Antioxidant Enzymes
All kinds of stress culminate in oxidative stress caused by reactive oxygen species (ROS) such as superoxide (O2•-) radicals, hydrogen peroxide (H2O2), and hydroxyl (OH•) radicals (Imlay 2003). Plants have evolved many protective mechanisms against ROS. These mechanisms include the production of antioxidants and antioxidative enzymes, for instance, catalases (CAT), superoxide dismutases (SOD), peroxidases (POD), and glutathione reductases (GR) (Ahire et al. 2012). Among various antioxidative enzymes, SOD, CAT, and POD make up the first line of defense in scavenging ROS. SOD converts superoxide radicals to H2O2, which is noxious to the nucleic acids, proteins, and chloroplast, and is dealt with by CAT and POD (Shen et al. 2010).
Si modulates the plant antioxidant defense system to prevent oxidative damage in the stressed plants (Kim et al. 2017). Several reports have described the Si-induced upregulation of antioxidative enzymes such as CAT, GR, SOD, guaiacol peroxidase (GPX), and ascorbate peroxidase (APX) (Shen et al. 2010; Soundararajan et al. 2014; Zhu and Gong 2014; Etesami and Jeong 2018), and peroxidase mediated host defense responses as well (Torres et al. 2006). Supplementation of Si had increased the level of POD in rice and cucumber plants challenged with Bipolaris oryzae and Podosphaera xantii (Dallagnol et al. 2011).
10 Si and Nutrient Uptake
The availability and uptake of nutrients in sufficient amounts is a prerequisite for healthy plant growth and architecture. Plant nutrients are primarily divided into two groups: macronutrients and micronutrients, based on the amount in which the plants require them. Any change in the optimum levels of any of these nutrients leads to abnormalities in plants (Shrivastav et al. 2020).
The application of Si is known to influence the uptake of macronutrients like N, P, and K in plants. Such an application elevated the level of N in cowpea (Mali 2008), wheat (Mali and Aery 2008), and rice (Singh et al. 2006; Detmann et al. 2012). The use of Si fertilizers increases the availability of P (Ma 2004; Singh et al. 2006) and influences the uptake of K (Kaya et al. 2006). In soybean, the application of Si was shown to improve the growth of plants and enhance the uptake of K (Miao et al. 2010). Si also mediates enhanced uptake of Ca and Mg (Kaya et al. 2006; Mali and Aery 2008). Moreover, the presence of Si not only reduces the uptake of heavy metals like Al and Cd (Ma and Takahashi 2002; Ma et al. 2004) but also mitigates the deficiency of micronutrients like Fe, Mn, Cu, Zn, and B in plants (Pavlovic et al. 2013; Hernandez-Apaolaza 2014).
11 Conclusions
Adverse environmental conditions adversely affect plant growth, development, and yield. Si plays a vital role in the alleviation of stress caused by various harsh environments. It influences multifunctional traits such as growth, morphology, the activity of antioxidant enzymes, accumulation of osmolytes, photosynthesis, and nutrient uptake in plants. The ability to take up Si under different environmental conditions varies from species to species. Moreover, the effects and their magnitudes caused due to Si supplementation vary from species to species and the prevalent conditions. Substantial evidence exists that underline the beneficial role of Si in plants under abiotic stress, but how Si manipulates the mechanisms of alleviation is still much of a mystery.
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Acknowledgments
Financial assistance by the University Grants Commission, New Delhi, and Department of Science and Technology (DST), Government of India and DST-PURSE are gratefully acknowledged. Authors are also grateful to Yashavantrao Chavan Institute of Science, Satara for financial support under DBT-STAR college scheme. Financial assistance to the faculty under self-funded project is acknowledged and also for providing facilities.
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Mundada, P.S. et al. (2021). Silicon and Plant Responses Under Adverse Environmental Conditions. In: Husen, A. (eds) Plant Performance Under Environmental Stress . Springer, Cham. https://doi.org/10.1007/978-3-030-78521-5_14
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