Silicon in agriculture: A review of its roles, mechanisms, and applications in crop production

Introduction

Approximately 28% of the Earth's crust consists of silicon (Si) by weight, and silicon is found in almost all agricultural soils, though it has been neglected in plant nutrition because it is not essential to the completion of the life cycle of the majority of plants (Epstein, 1999; Datnoff et al., 2001) [2,1] . Consequently, silicon has always been considered a "beneficial" or "quasi-essential" element rather than an essential one (Elsokkary, 2018) [3] , since while the lack of silicon in soils does not affect the process of growth, its presence increases the performance of plants especially when they are exposed to various stresses (Richmond et al., 2003; Guntzer et al., 2012; Coskun et al., 2019) [5,6,4] . At the same time, the interest towards silicon has been rising since, owing to intensive cropping practices, harvesting cycles, and the usage of synthetic fertilizers poor in silicon, the availability of silicon for plants has been declining (Savant et al., 1996; Sommer et al., 2006) [7,8] , which has occurred during a period marked by climatic changes, soil salinity, and pests. Interest in this element was not generated recently. Japanese plant pathologists observed the relationship between the level of silicon in the soil and rice blast disease at the beginning of the twentieth century (Pooja et al., 2014) [9] . Over many years of soil fertility and plant physiology studies, researchers have shown the need to pay attention to silicon because of its importance for agronomy (Fox et al., 1978; Marschner, 1995; Ma, 2004) [10,11,12] . Identification of the special gene that encodes silicon transporters at the beginning of the 2000s (Ma et al., 2006; Ma et al., 2007) [13,14] provided the molecular base to this research field. Since then, there has been a rapid increase in the number of publications related to soil science, plant physiology, molecular biology, entomology, plant pathology, and nanotechnology (Frew et al., 2018; Coskun et al., 2019) [15,4] . This review highlights current research studies and reviews involving soil silicon chemistry, silicon absorption and transportation by plants, the effect of silicon on alleviation of abiotic and biotic stresses, its impact on crop productivity and quality in various plant species, and the novel field of silicon nanoparticles along with their effects on rhizospheric microbiome. All these pieces of evidence together help establish a clear trend regarding the fact that silicon needs to be incorporated more systematically into soil fertility management practices (Guntzer et al., 2012; Meharg et al., 2015; Debona et al., 2017) [6,17,16] .

Silicon in soil: Forms, sources, and availability

Silicon in soil is found in three forms-liquid, adsorbed, and solid (Sommer et al., 2006) [8] . Silicon in its plant-available form, monosilicic acid (H 4 SiO 4 ), is soluble in soil under pH conditions less than 9, and its concentration in soil solution is usually comparable to other elements, like potassium and sulfate (Epstein, 1999; Ma et al., 2006) [2,18] . Mineral soils have a high total amount of silicon in the form of quartz sand and primary silicates (olivine, feldspar, mica) and also secondary clay silicates (illite, kaolinite, montmorillonite), but since the total amount is insoluble and biogeochemically inactive, it is not possible to use total soil silicon to predict the amount available to plants (Sommer et al., 2006; Cornelis et al., 2016) [8,19] . The availability of silicon is low in highly weathered and leached soils, like Oxisols and Ultisols, in organic Histosols that have little mineral content, and in soils that have high quartz sand content or a long history of intense cropping in the absence of silicon supply (Savant et al., 1996; Klotzbücher et al., 2015) [21,20,7] . Soil pH, soil redox potential, soil organic matter, and soil oxides of iron and aluminum all affect the amount of silicic acid left in soil solution as opposed to the amount of silicic acid adsorbed onto the mineral surface (Sommer et al., 2006; Gérard et al., 2008) [8,22] . Soil waterlogging increases solubility of silicon, while adsorption on iron and aluminum hydroxides decreases the solubility of silicic acid in the soil solution as the soil pH increases (Cornelis et al., 2016) [19] . This information is important for agriculture since the actual threshold for silicon fertilizer application is not the total silicon content of the soil but rather its plant-available content, measured with extractants like citric acid, acetic acid, or sodium acetate (Savant et al., 1996; Klotzbücher et al., 2015 [7] ; Heckman, 2021). There is a biological cycling mechanism whereby silicon moves back to the soil in large part due to plants themselves. In a mechanism whereby plants that accumulate silicon take in monosilicic acid, the process of polymerization in the plants leads to silica bodies known as phytoliths, which are amorphous and hydrated (Currie et al., 2007; Trembath-Reichert et al., 2015) [23,24] . When the plant dies and its tissues decay, the phytoliths get dissolved and hence contribute to the pool of available silicon in the soil (Strömberg et al., 2018) [25] . It follows that residue management such as recycling rice straw and husk would influence the soil's availability of silicon in the long term (de Tombeur et al., 2021; Cornelis et al., 2016) [26,19] . [18,23] . Silicon once deposited is immobile and cannot be translocated to younger parts (Ma et al., 2008) [27] ; thus, its frequent application gives better results than once-off application. There are three ways of silicon uptake: active silicon uptake where the roots absorb silicon at a higher rate compared to water, hence creating low concentrations of silicon around them; passive uptake, whereby uptake takes place in proportion to the transpiration flow; and rejective uptake where plants exclude silicon. There is great variation among species as far as the uptake of silicon goes and the amounts of silicon accumulated. From the concentrations of silicon in the shoots and their ratios to calcium, plants can be categorized into accumulators (silicon content above 1%), intermediate accumulators, and excluders (less than 0.5%). Accumulators include rice, sugarcane, and many members of the Poaceae and Cyperaceae families. The molecular basis for this variation became clear with the discovery of dedicated silicon transporter genes. Transporter genes for influx (type Lsi 1 , low-silicon genes which belong to the nodulin26-like intrinsic proteins group) transport silicic acid from the soil solution to root cells, whereas efflux transporter genes (type Lsi 2 ) transport it from root cells to the xylem for delivery to the shoots; there is an ad-ditional transporter gene (type Lsi 6 ) involved in silicon unloading from the xylem to leaf tissue (Ma et al., 2006; Ma et al., 2007; Yamaji et al., 2007; Yamaji et al., 2011) [13,14,28,29] . The transporters were initially identified in rice but have later been found in barley (Chiba et al., 2009) [30] , maize (Mitani et al., 2009) [31] , wheat (Montpetit et al., 2012) [32] , soybean (Deshmukh et al., 2013) [33] , cucumber, and other plants, giving an insight into the genetic basis behind the different amounts of silicon accumulated by various crops growing in the same soils (Ma et al., 2015; Kaur et al., 2019) [35,34] .

Uptake and transport of silicon in plants

Silicon and abiotic stress tolerance

Drought and water-deficit stress

The application of silicon, either as soil application, foliar spray or nanosilicon, has time and again proven effective on improving growth and development in crops such as wheat, sugarcane, sugar beet, maize, soybean and other crops, under water stress conditions (Wang et al., 2021; Zhu et al., 2014) [36,37] . Mechanisms reported include improved cell membrane integrity and relative water content, higher leaf chlorophyll content and photosynthesis rate, higher root hydraulic conductivity and radial water transport, reduced water loss by transpiration due to formation of silica in leaves, and greater antioxidant enzyme activities including superoxide dismutase, catalase, ascorbate peroxidase which prevent oxidation stress due to the generation of reactive oxygen species due to drought stress (Kim et al., 2017; Wang et al., 2021; Biju et al., 2017) [39,36,38] . In lentils, foliar and root-applied silicon regulates osmolyte and hydrolytic enzymes activity and reprogramming gene expression pathways for drought stress tolerance (Biju et al., 2017; Biju et al., 2021; Biju et al., 2023) [38,40,41] . In rice, silicon fertilization enhanced plant water balance, photosynthesis rate, and uptake of minerals under water deficit condition (Chen et al., 2011) [42] , while the combination of water management practices and silicon treatments resulted in decreased accumulation of arsenic as well as in improved germination and growth (Zia et al., 2017) [43] . For the sugarcane crop, silicon fertilization increased photosynthetic efficiency and activity of antioxidant enzymes under conditions of drought (Li, 2019) [44] . In a number of field and pot experiments, silicon-fertilized plants subjected to water deficit exhibited higher yields, biomass, and grain quality compared to non-treated plants, and especially under the conditions of more intense drought (Zhu et al., 2014; Debona et al., 2017) [37,16] .

Salinity stress

Silicon acts against salinity stress using a variety of strategies: lowering the uptake and transport of sodium into shoots, increasing the absorption of potassium, thereby raising the K + /Na + ratio, ion compartmentalization into the vacuoles, lowering membrane permeability and electrolyte leakage, and increasing the activities of antioxidative enzymes (Liang, 1999 [46,45,37] . Silicon treatment in barley plants led to changes in the ultrastructure of leaves, chlorophyll concentration and photosynthesis in saltstressed plants (Liang, 1998) [47] , and hydroponic silicon application in soybean caused enhanced growth under salinity stress (Lee et al., 2010) [48] . The same has been reported for cucumber, tomato, sweet pepper, spinach, and cereal crops under saline conditions, leading to better germination, growth, and biomass production (Abdelaal et al., 2020; Dhiman et al., 2021) [49,45] .

Heavy-metal and mineral toxicity

Another common feature of the reviewed literature is that silicon can decrease the uptake, translocation, and toxicity of aluminum, cadmium, arsenic, manganese, lead, zinc, and many more harmful elements (Neumann & zur Nieden, 2001; Wu et al., 2013) [50,51] . The following mechanisms have been suggested to account for these effects: precipitation of metals outside the cells along with silica precipitation in the growth medium, changes in the rhizosphere pH, increased by silicon, reducing the solubility of the metals, metal binding to the cell walls and their limitation in entering the cytoplasm, hydroxy-aluminosilicate complexation as a mechanism of aluminum detoxification, and enhanced activities of antioxidative defense systems against metal-induced oxidative stress (Hou et al., 2023; Liang et al., 2007) [52,53] . For example, it was found that silicon nutrition decreased cadmium uptake and reprogramming of leaf proteome during cadmium stress in rice (Nwugo et al., 2008; Nwugo et al., 2011) [ 54, 55] and that silicon nutrition decreased metal accumulation and translocation but increased plant biomass and chlorophyll concentration in the hyperaccumulator of arsenic/cadmium Isatis cappadocica (Azam et al., 2021) [56] . Zinc toxicity reduction due to zinc transport regulation and antioxidant activity has also been reported in rice (Song et al., 2011; Gu et al., 2012) [58,57] . Such effects have great significance in crop production in acidic, metal-polluted, or heavily weathered soils in which aluminum or manganese toxicity is an obstacle for growth (Wu et al., 2013; Zargar et al., 2019) [51,59] .

Other abiotic stresses

Silicon has been found to reduce the impact of extreme temperatures, UV-B exposure, nutritional stress, and lodging (Etesami et al., 2018; Bakhat et al., 2018) [61,60] . In various studies involving cereals, the presence of silicon within stem tissues resulted in structural hardness and lignification, thus preventing lodging and preserving canopy photosynthesis and productivity during wind and rain (Detmann et al., 2012) [62] . Silicon has also been shown to enhance phosphorus uptake and utilization efficiency, possibly because of its influence on the secretion of organic acids and phosphate transporters from root tissues (de Tombeur et al., 2021) [26] .

Silicon and biotic stress management

Plant pathogens

Silicon is arguably one of the most reliable methods of decreasing the impact of both fungi and bacterial diseases on various crops (Fauteux et al., 2005; Wang et al., 2017; Rodrigues, 2015) [63,64,65] . There are two basic mechanisms that are thought to be involved. The first one is physical. Silicon polymerization under the leaf cuticle results in formation of a physical barrier-the so-called double cuticular layer, which prevents fungi from penetrating into plant tissues and also silica deposition at the infection points (papillae) creates a physical barrier against pathogens (Rodrigues et al., 2003; Guerriero et al., 2016) [67, 66] . The second mechanism is biochemical or molecular. It has been shown that silicon application enhances the activity of certain defense enzymes such as peroxidase, polyphenol oxidase, phenylalanine ammonia-lyase, and chitinase (Cai et al., 2008; Mohammadi et al., 2002) [68,69] , increases the concentration of antimicrobial phenols and phytoalexins (Fawe et al., 1998; Rodrigues et al., 2004; Remus-Borel et al., 2005) [70,71,72] , and primes plants for enhanced reaction to pathogens by promoting gene expression (priming effect) (Vivancos et al., 2015; Van Bockhaven et al., 2013) [74,73] . Numerous studies have shown that the use of silicon is highly effective in inhibiting the intensity of pathogens affecting crop species. Silicon applications on cereals inhibit the effect of the major fungal diseases such as blast, brown spot, and sheath blight in rice (Rodrigues et [67,76,75] , while on wheat and barley it prevents the development of powdery mildew and spot blotch (Carver et al., 1987; Bélanger et al., 2003; Perez et al., 2014) [78,77,79] . The same effects are recorded in perennial and horticulture crops such as rust and ring spot on sugarcane, Panama wilt and black Sigatoka on banana, leaf rust in coffee (Carré-Missio et al., 2009) [80] , powdery mildew in cucurbits (Fawe et al., 1998) [70] and grapevine (Bowen et al., 1992) [81]

, bacterial wilt and speck in solanaceous crops such as tomato and potato (Andrade et al., 2013) [82] . Despite the fact that silicon applications do not inhibit the initial penetration of pathogens into the plants, they significantly reduce the development of the disease leading to the smaller lesions.

Arthropod pests and herbivores

The use of silicon causes plants to gain resistance against different insect herbivores and mites using a combination of physical and biochemical processes (Reynolds et al., 2016; Alhousari et al., 2018) [84,83] . Physically, deposition of silicon makes the leaves more abrasive and rigid; as a result, palatability, digestibility, and mechanical wear of herbivore mouthparts are reduced in some cases (Massey et al., 2006; Massey et al., 2009; Keeping et al., 2008) [85,86,87] . Biochemically, silicon plays an active role in jasmonic acid signaling process that is involved in inducible defenses against chewing herbivores. Some studies demonstrate the presence of positive feedback between jasmonate signaling and silicon accumulation (Ye et al., 2013; Reynolds et al., 2016) [88,84] . Silicon has similarly been implicated in alterations in the volatile emissions of the HIPVs which could enhance the attraction of their enemies such as predators and parasitoids towards the stressed plants. This is yet another type of indirect defense mechanism of the third trophic level aside from the direct one exhibited by the herbivores (Reynolds et al., 2016; Golubkina et al., 2022) [84,89] . Its effects have been reported for leaffolder insect pest of the rice (Ye et al., 2013) [88] and diverse chewing and piercing-sucking pests on both grasses and legumes, as well as the vertebrate herbivores in some grass and cereals (Massey et al., 2006; Alhousari et al., 2018) [87,83] .

Silicon fertilization: Effects on crop yield and quality

Trials on cereals, legumes, root and tuber crops, vegetables, fruit and ornamental plants consistently show that application of silicon fertilizers results in increased yields or improvement in quality, either via soil application or through foliar spraying (Artyszak, 2018; Sarma et al., 2024) [90,91] . Since silicon is not retranslocated once it is absorbed, any positive effects can be observed only under some form of stress, whether drought, salinity, disease or pest attack, and foliar application of nutrients has been found relatively inexpensive and easy compared to soil applications (Artyszak, 2018; Munir et al., 2023) [90,92] .

Cereals

In wheat, silicon fertilization-via sodium silicate, silicate slag, potassium metasilicate, or proprietary silicic acid formulations-has increased grain yield primarily by improving spike density, number of grains per spike, and thousand-grain weight, with particularly strong effects reported under drought and salinity stress and in soils with low native silicon availability (Artyszak, 2018; Dhiman et al., 2021) [90,45] . Similarly in maize, increased yield was achieved due to increased number of grains per ear and grain weight, as well as better drought tolerance (Sarma et al., 2024; Munir et al., 2023) [91,92] . In rice, silicon fertilization has increased grain yield through improved spikelet formation, fertilization, and grain filling (Detmann et al., 2012; Klotzbücher et al., 2015) [62,20] , alongside its well-established role in disease suppression (Rodrigues et al., 2003; Datnoff et al., 2001) [67,1] .

Legumes and oilseeds

In soybean, foliar silicic acid has increased the number of pods per plant, seeds per pod, and thousand-seed weight, with benefits most evident under drought and salinity stress (Lee et al., 2010; Artyszak, 2018) [48,90] . In lentil, silicon supplementation improved seed germination, osmotic regulation, and antioxidant defense under water-limited conditions (Biju et al., 2017; Biju et al., 2021) [38,40] . In winter oilseed rape, silicon-containing foliar products have improved winter survival via more compact, cold-hardy rosette growth, root system development, and seed weight (Artyszak, 2018) [90] .

Root and tuber crops

Sugar beet trials, particularly from European field studies, report consistent increases in root yield, biological and commercial sugar yield, and in some cases a reduction in impurities such as alpha-amino nitro gen that lower processing quality, following silicon fertilization delivered as marine calcite, potassium or sodium silicate solutions, or nano-silica formulations (Artyszak et [93] ; Artyszak & Gozdowski, 2021; Viciedo et al., 2019) [94] . In potato, silicon fertilization has increased average tuber weight and total tuber yield, altered tuber skin anatomy in ways associated with delayed maturation and improved storage, reduced nitrate accumulation, and in some studies reduced the incidence of specific tuber diseases, although effects on overall disease pressure have been inconsistent across studies (Trawczyński, 2018; Artyszak, 2018) [95, 90] .

Vegetables, berries, and orchard crops

Across lettuce, tomato, cucumber, pepper, onion, and strawberry, silicon fertilization-soil or foliar-has generally increased yield, fruit or head weight, firmness, and postharvest quality, while in several cases reducing fruit cracking, nitrate accumulation, and susceptibility to specific pathogens (Peris-Felipo et al., 2020; Hajiboland et al., 2018; Abdelaal et al., 2020) [96,97,49] . In orchard and vine crops, silicon applications have reduced grapevine powdery mildew severity (Bowen et al., 1992) [81] , increased grape yield, and improved seedling establishment in ornamental bedding and pot plants through firmer, better-formed growth (Artyszak, 2018) [90] .

Commercial silicon fertilizer sources

Silicon fertilizers used in practice include solid mineral sources such as calcium silicate, potassium silicate, sodium silicate, and calciummagnesium silicate-often applied to soil and, in the case of calcium silicate, valued additionally as a liming material; liquid stabilized silicic acid formulations used mainly for foliar application; and industrial by-products such as iron and steel slag, blast-furnace slag, and fly ash, which supply silicon while also recycling waste material (Datnoff et al., 2001; Artyszak, 2018; Munir et al., 2023) [1,90,92] . Plant-derived silica sources, including rice hull and rice hull ash, offer an additional, renewable route to replenishing plant-available soil silicon in intensively cropped systems such as rice paddies (Klotzbücher et al., 2015; Savant et al., 1996; Pereira et al., 2004) [20,7,98] , and silicon is now recognized in several countries as an officially designated plant-beneficial substance eligible for fertilizer labeling (Heckman, 2013) [99] .

Silicon-based nanoparticles in agriculture

A growing body of work examines silicon dioxide nanoparticles (SiO 2 -NPs or Si-NPs) as a higher-bioavailability alternative to conventional silicate fertilizers, which are often poorly soluble (Rajput et al., 2021) [100] . It has been observed that Si-NPs can help enhance germination and vigor of plants including wheat, peas, mustard, sunflowers, tomatoes, lentils, and maize, increase the content of photosynthetic pigments and gas exchange capacity, and improve resistance against salinity, drought, low temperatures, and UV-B through a mechanism very much like that in bulk silicon but with higher efficiency at smaller quantities due to the large surface-to-volume ratio and activity of nanoparticles (Sharf-Eldin et al., 2023; Rajput et al., 2021) [101,100] . Si-NPs have also shown antimicrobial activity against a range of plant pathogens and can reduce the uptake and phytotoxicity of heavy metals such as cadmium, lead, arsenic, and mercury by limiting their translocation from root to shoot and by supporting antioxidant defense systems (Rajput et al., 2021; Hou et al., 2023) [100,52] . Mesoporous silica nanoparticles have additionally been explored as carriers for macronutrients and pesticides in controlled-release fertilizer and nanopesticide formulations, exploiting their large surface area, tunable pore size, and stimulus-responsive release behavior to improve the efficiency and photostability of agrochemicals while reducing environmental losses. As with conventional silicon, the magnitude and direction of Si-NP effects depend heavily on particle size, shape, concentration, and method of application (soil drench, seed priming, or foliar spray), and soil-applied Si-NPs are generally re-ported to be more effective than foliar applications for producing measurable increases in plant tissue silicon content (Rajput et al., 2021) [100] .

Silicon, soil properties, and the rhizosphere microbiome

In addition to their direct impacts on the growth of plants, silicon and Si-NPs affect the soil-plant-microbe ecosystem. It has been demonstrated that amendments of silicon have an effect on the metabolic composition of the rhizosphere-the presence of organic acids, sugars, and fatty acids related to carbon and nitrogen metabolism, altering the ratio of certain genera of bacteria and fungi in the rhizosphere, increasing the growth-promoting genera of bacteria, such as Paenibacillus, as well as interacting with phosphate-solubilizing and nitrogen-fixing microorganisms, thus positively affecting nutrient cycling and soil fertility (Rajput et al., 2021; Cornelis et al., 2016) [100,19] . In turn, silicate-solubilizing bacteria, which are found in the soil, are capable of transforming insoluble mineral silicates into soluble forms that can be assimilated by the plants. This process provides a biological alternative to silicon fertilization (Rajput et al., 2021; Etesami et al., 2018) [100,61] . However, there have been some reports on the negative impact of excessive amounts of Si-NPs on the activity of soil enzymes and microbial gene abundance (Rajput et al., 2021; Wu et al., 2013) [100,51] .

Conclusion

Silicon is increasingly recognized as an important beneficial element in modern crop production, particularly under conditions of environmental and biological stress. While silicon is not classified as one of the elements that are essential for the entire life cycle of the plant, the information that has been provided above clearly indicates that silicon can bring a lot of benefit to plant growth, productivity and resistance through its impact on soil-plant interactions. The benefits that silicon offers to plants come through the complex dual interaction that is composed of two aspects -the physical and the biochemical aspects. This combination enables plants to cope up with various stressful situations including droughts, salinity, heavy metal toxicity, fungi attacks, pests and others. Despite the advances that have been made in this area, many knowledge gaps still exist. There is the possibility that plant-available silicon could be reduced as a result of intensive farming practices, continuous removal of residue, and the use of silicon-deficient synthetic fertilizers; but silicon management has received limited attention within soil testing and fertilizer recommendations. The recent developments in silicon fertilization and crop residue management, as well as the novel techniques of using silicon nanoparticles (Si-NPs) and silicate-solubilizing bacteria, hold great promise in this regard. However, there is a lack of knowledge regarding the efficiency of these methods on the farm-scale, optimal application rates, and environmental fate of silicon-based nanomaterials.

In general, the scientific literature reviewed in this paper shows that the use of silicon as a fertilizer input can be included within broader frameworks of soil fertility and crop climate resilience management, especially for high-input and stress-prone agroecosystems. Silicon should not be seen as another fertilizer input but rather as an integral part of integrated soil-plant-microbe management system.

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