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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Plant Sci.</journal-id>
<journal-title>Frontiers in Plant Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Plant Sci.</abbrev-journal-title>
<issn pub-type="epub">1664-462X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fpls.2025.1641798</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Plant Science</subject>
<subj-group>
<subject>Hypothesis and Theory</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>The positive impact of silicon on the yield and quality of tobacco</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>He</surname>
<given-names>Mengjie</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/3146473/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/conceptualization/"/>
<role content-type="https://credit.niso.org/contributor-roles/data-curation/"/>
<role content-type="https://credit.niso.org/contributor-roles/software/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Li</surname>
<given-names>Qiang</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/3032315/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/funding-acquisition/"/>
<role content-type="https://credit.niso.org/contributor-roles/project-administration/"/>
<role content-type="https://credit.niso.org/contributor-roles/resources/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Ma</surname>
<given-names>Ya</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/formal-analysis/"/>
<role content-type="https://credit.niso.org/contributor-roles/supervision/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Zhou</surname>
<given-names>Pengtao</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/conceptualization/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Kang</surname>
<given-names>Kang</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/formal-analysis/"/>
<role content-type="https://credit.niso.org/contributor-roles/supervision/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Wu</surname>
<given-names>Boran</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/conceptualization/"/>
<role content-type="https://credit.niso.org/contributor-roles/investigation/"/>
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</contrib>
</contrib-group>
<aff id="aff1">
<sup>1</sup>
<institution>College of Resources, Hunan Agricultural University</institution>, <addr-line>Changsha, Hunan</addr-line>,&#xa0;<country>China</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>College of Agriculture, Hunan Agricultural University</institution>, <addr-line>Changsha, Hunan</addr-line>,&#xa0;<country>China</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Alexander Lux, Comenius University, Slovakia</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Krishan K. Verma, Guangxi Academy of Agricultural Sciences, China</p>
<p>Arkadiusz Artyszak, Warsaw University of Life Sciences, Poland</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Qiang Li, <email xlink:href="mailto:zqiangli@126.com">zqiangli@126.com</email>
</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>04</day>
<month>08</month>
<year>2025</year>
</pub-date>
<pub-date pub-type="collection">
<year>2025</year>
</pub-date>
<volume>16</volume>
<elocation-id>1641798</elocation-id>
<history>
<date date-type="received">
<day>05</day>
<month>06</month>
<year>2025</year>
</date>
<date date-type="accepted">
<day>14</day>
<month>07</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2025 He, Li, Ma, Zhou, Kang and Wu.</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>He, Li, Ma, Zhou, Kang and Wu</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>Silicon (Si) is widely used in agricultural crop practices. However, the effects of varying Si application rates on tobacco growth and quality remain unclear. Therefore, this study applied four different Si concentrations, i.e., 0, 750, 1500 and 3000 kg/ha of Si (S0, S50, S100 and S200), examined the impact of different Si concentrations on tobacco (<italic>Nicotiana tabacum</italic> L. &#x2018;Yunyan 87&#x2019;) growth, nutrient utilization, and economic quality under field conditions. The results demonstrated that Si application significantly improved tobacco growth, the biomass significantly increased by 19.5%-26.53%; during button stage, the plant height significantly increased by 15.38%-19%. Si also enhanced nutrient use efficiency, particularly for nitrogen and potassium. The utilization efficiency of N and K fertilizer were significantly increased by 27.42%-43.71% and 40.25% - 44.63%, respectively. Furthermore, Si improved leaf physical properties, enhancing single-leaf weight and leaf area, while reducing leaf density and midrib ratio, optimizing leaf quality by improving the sugar-alkali ratio and potassium-chloride balance. Notably, the reducing sugar content in upper leaves increased by 15.21% with S50 treatment, while the chlorine content in middle leaves was decreased by 11.11% with S100. Additionally, among all treatments, S50 achieved the highest proportion (94.75%) of medium and high-quality tobacco leaves, along with a 15.70% increase in yield and a 30.76%&#xa0;boost in output value compared to S0. However, excessive Si&#xa0;application (3000 kg/ha) negatively affected quality, increasing nicotine levels and disrupting the sugar-alkali ratio, which elevate leaf irritancy. In conclusion, moderate Si application (750&#x2013;1500 kg/ha) is an effective strategy for enhancing tobacco yield and quality, offering a sustainable approach to optimize cultivation practices.</p>
</abstract>
<kwd-group>
<kwd>silicon</kwd>
<kwd>tobacco</kwd>
<kwd>growth</kwd>
<kwd>nutrient use efficiency</kwd>
<kwd>economic benefits</kwd>
</kwd-group>
<contract-sponsor id="cn001">Hunan Normal University<named-content content-type="fundref-id">10.13039/501100002664</named-content>
</contract-sponsor>
<counts>
<fig-count count="4"/>
<table-count count="6"/>
<equation-count count="7"/>
<ref-count count="46"/>
<page-count count="14"/>
<word-count count="8412"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Plant Nutrition</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>Tobacco (<italic>Nicotiana tabacum</italic> L.) as a widely cultivated economic crop, is extensively grown in Brazil, China, the United States, and India (<xref ref-type="bibr" rid="B23">Lisuma et&#xa0;al., 2023</xref>). As a globally traded crop, tobacco yields and quality is an important indicator of tobacco plant that determines farmers&#x2019; income and tobacco market prices (<xref ref-type="bibr" rid="B42">Yan et&#xa0;al., 2025</xref>). Therefore, the yield and quality of tobacco leaves deserve special attention.</p>
<p>Silicon (Si), as a beneficial element for plants, is widely used in rice, tomatoes, sugarcane and other high Si demanding crops (<xref ref-type="bibr" rid="B43">Yan et&#xa0;al., 2023</xref>; <xref ref-type="bibr" rid="B11">Fan et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B38">Verma et&#xa0;al., 2020a</xref>). Researchers found that Si promoted the root development of plants, increased photosynthetic efficiency and the dry matter mass, thereby increased yields (<xref ref-type="bibr" rid="B18">Ji et&#xa0;al., 2017</xref>). Besides, Si improved the stress resistance of plants, such as salt stress, heavy metal toxicity, drought, high temperature, and ultraviolet radiation (<xref ref-type="bibr" rid="B39">Verma et&#xa0;al., 2023</xref>; <xref ref-type="bibr" rid="B38">Verma et&#xa0;al., 2020a</xref>; <xref ref-type="bibr" rid="B12">Ghosh et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B9">El-Saadony et&#xa0;al., 2022</xref>). And Si effectively reduced crop lodging and improved insect resistance by enhancing the mechanical strength such as stems and cell walls, especially under adverse weather conditions (<xref ref-type="bibr" rid="B20">Kumari et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B19">Khan et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B30">Pavlovic et&#xa0;al., 2021</xref>). In addition, the application of Si not only impacted plants, but also had a significant impact on soil physicochemical properties and microbial community structure (<xref ref-type="bibr" rid="B2">Artyszak, 2018</xref>). Si application altered the chemical composition of soil minerals by promoting the formation of new clay minerals (<xref ref-type="bibr" rid="B36">Thakral et&#xa0;al., 2024</xref>). These clay minerals exhibited a high specific surface area, enabling the adsorption of water, nutrients nitrogen (N), phosphate (P), potassium (K), aluminum (Al), and heavy metals through both physical and chemical mechanisms, which increased soil available Si content, reduced soil heavy metal toxicity, regulated soil pH value, enhanced soil water retention and permeability, thereby improved soil quality, and promoted nutrient use efficiency (<xref ref-type="bibr" rid="B35">Souri et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B34">Song et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B33">Schaller et&#xa0;al., 2021</xref>).</p>
<p>However, it was reported that excessive application of Si may have negative effects, which lower soil pH, leading to soil acidification, and affected soil microbial activity (<xref ref-type="bibr" rid="B33">Schaller et&#xa0;al., 2021</xref>). Besides, Si competes with other nutrients such as N, P and K for absorption, and excessive application may lead to nutrient imbalance, affecting normal plant growth (<xref ref-type="bibr" rid="B7">Coskun et&#xa0;al., 2019</xref>). Researchers have found that, the demand for Si varies among different plants. For example, rice is a high Si demanding crop with a critical concentration of 30-50g/kg, while the Si critical concentration of tomatoes is 5-8g/kg (<xref ref-type="bibr" rid="B36">Thakral et&#xa0;al., 2024</xref>).</p>
<p>In addition, current researches on Si fertilizer predominantly focuses on field crops harvested for seeds and fruits, as well as&#xa0;vegetables, whereas tobacco cultivation primarily targets leaf&#xa0;production (<xref ref-type="bibr" rid="B36">Thakral et&#xa0;al., 2024</xref>). However, there is limited research on the effects of Si fertilizer on tobacco. The appropriate application amount of Si fertilizer on tobacco is not clear, and more research is needed to explore the appropriate application amount of Si fertilizer in tobacco agricultural production. Based&#xa0;on this, we conducted field experiments on <italic>Nicotiana tabacum</italic> L. &#x2018;Yunyan 87&#x2019;, to investigate the influence of different rate of Si fertilizer on tobacco growth, yields and quality, five experimental treatments were established in this study: CK (without fertilization), S0 (control; conventional fertilization), S50 (application of Si fertilizer at 750 kg/ha), S100 (application of Si fertilizer at 1500 kg/ha), and S200 (application of Si fertilizer at 3000 kg/ha).The agronomic traits, physical characteristics, quality characteristics, yield, output value, average price, nutrient accumulation and fertilizer utilization rate of N, P, K, were assayed in this study. This study to provide a scientific basis for rational Si fertilizer utilization in tobacco cultivation, and practical fertilizer usage guidelines for tobacco farmers to improve tobacco quality and overall market value.</p>
</sec>
<sec id="s2" sec-type="materials|methods">
<label>2</label>
<title>Materials and methods</title>
<sec id="s2_1">
<label>2.1</label>
<title>Experiment design and field location</title>
<p>The tobacco variety (<italic>Nicotiana tabacum</italic> L. &#x2018;Yunyan 87&#x2019;) was used in this study, which has good disease resistance, coordinated dehydration and drying, and is easy to bake. The field experiment was conducted in Lanshan County, Yongzhou City, Hunan Province, China (25.18&#xb0;N, 112.18&#xb0;E) from March to August 2024. The specific location is shown in <xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>. The annual average temperature in this area is 18&#xb0;C to 19 &#xb0;C, the annual precipitation is 1400 to 1600 millimeters, and the annual sunshine hours are 1400 to 1600 hours. The climate data from March to August, including temperature, precipitation and hydrothermal index are shown in <xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>. The basic physicochemical properties and the microelement content of the soil (0&#x2013;20 cm) were as follows: pH, 6.73; organic matter (OM), 65.97 mg/kg; alkali-hydrolysable nitrogen (AN), 229.10 mg/kg; available phosphorus (AP), 102.81 mg/kg; available potassium (AK), 355.85 mg/kg; total nitrogen (TN), 3.64 g/kg; total phosphorus (TP), 2.42 g/kg; total potassium (TK), 6.63 g/kg; chloride ion (Cl<sup>-</sup>), 5.74 mg/kg; boron (B), 3.69 (mg/kg); available SiO<sub>2</sub>, 6.98 mg/kg; manganese (Mn), 38.92 mg/kg; iron (Fe), 357.89 mg/kg; nickel (Ni), 3.21mg/kg; copper (Cu), 2.08 mg/kg; zinc (Zn), 9.14 mg/kg.</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>The experimental site location. The pentagram mark in the picture indicates the location of the experimental field.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1641798-g001.tif">
<alt-text content-type="machine-generated">Map showing the location of Lanshan in Hunan Province within China. The main map highlights China in red, with Hunan in light green. Enlarged sections highlight Hunan and Lanshan in detail, with arrows indicating their positions.</alt-text>
</graphic>
</fig>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Climate data from March to August. On the left are precipitation and hydrothermal index, and on the right are temperature.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1641798-g002.tif">
<alt-text content-type="machine-generated">Bar and line graph showing precipitation, hydrothermal index, and temperature from March to August. Precipitation is highest in April and June, while the temperature steadily increases, peaking in July, then slightly decreases in August.</alt-text>
</graphic>
</fig>
<p>Tobacco was transplanted on March 27, 2024, the lower leaves were harvested on June 26, 2024, the middle leaves were harvested on July 15, 2024, and the upper leaves were harvested on August 1, 2024. The apical part was excised 2&#x2013;3 weeks prior to floral anthesis, and basal leaf removal was implemented when 30&#x2013;50% of the central inflorescence had reached bloom stage, ultimately retaining 18 functional leaves per plant through systematic defoliation. In order to ensure the growth and development of plants, high ridge cultivation was adopted during transplantation, covered with plastic film. During the period of plant growth, leaf diseases and leaf eating pests were controlled, and solar insecticidal lamps were installed in the field.</p>
<p>Five experimental treatments were established, CK is no fertilization, S0 is normal fertilization, S50 is normal fertilization+750 kg/ha Si fertilizer, S100 is normal fertilization+1500 kg/ha Si fertilizer and S200 is normal fertilization+3000 kg/ha Si fertilizer. The plant spacing and row spacing are 50 &#xd7; 120&#xa0;cm. Three replicates were set up for each treatment, with each replicate area of 4&#xa0;m &#xd7; 8.4&#xa0;m, and each treatment was randomly distributed.</p>
<p>The Si fertilizer used contains SiO<sub>2</sub> 50%, CaO 12%, MgO 0.7%, Fe<sub>2</sub>O<sub>3</sub> 2.3%, organic matter 10%, humic acid 6% and other trace elements 3%. Fertilizer application refers to <xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>. The nutrient input of each treatment was 185.4 kg/ha of N, 127.5 kg/ha of P<sub>2</sub>O<sub>5</sub>, and 430.5 kg/ha of K<sub>2</sub>O.</p>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Fertilizer application.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="center">Time</th>
<th valign="middle" align="center">Fertilization type</th>
<th valign="middle" align="center">Application ratio</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">5 days before  transplanting</td>
<td valign="middle" align="center" style="background-color:#ffffff">Base Fertilizer (1st)</td>
<td valign="middle" align="center" style="background-color:#ffffff">70%</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">3 days after transplanting</td>
<td valign="middle" align="center" style="background-color:#ffffff">Base Fertilizer (2nd)</td>
<td valign="middle" align="center" style="background-color:#ffffff">30%</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">25 days after transplanting</td>
<td valign="middle" align="center" style="background-color:#ffffff">Topdressing</td>
<td valign="middle" align="center" style="background-color:#ffffff">Conventional fertilization</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>Fertilizer ingredients: tobacco specific base fertilizer (8-10-11); seedling fertilizer (20-9-0); tobacco specific topdressing (10-0-32); potassium sulfate (K<sub>2</sub>O=52.0%, Cl=1.5%, S=17.0%); silicon fertilizer (SiO<sub>2</sub> = 50%).</p>
</fn>
</table-wrap-foot>
</table-wrap>
</sec>
<sec id="s2_2">
<label>2.2</label>
<title>Collection of tobacco and soil samples</title>
<p>Soil samples ranging from 0&#x2013;20 cm were gathered from 10 random sites within each plot on March 15, 2024 and after visible plant residues and stones were removed. The soil samples were air-dried, pulverized, and sieved through 2&#xa0;mm and 0.0149&#xa0;mm screens for subsequent analysis, including pH, OM, HN, AP, AK, TN, TP, TK, Cl<sup>-</sup>, B, available SiO<sub>2</sub>, Mn, Fe, Ni, Cu, Zn.</p>
<p>At the flower budding stage and button stage of tobacco, 10 plants with consistent growth and no pests or diseases were randomly selected from each plot. The 3rd, 6th, and 9th leaves were selected from top to bottom for SPAD value measurement. Agronomic traits such as plant height (cm), stem circumference (cm), effective leaves (leaves/plant), maximum leaf length (cm), maximum leaf width (cm), and maximum leaf area (cm&#xb2;) were recorded for each treatment. At the maturating stage of tobacco, the harvested mature tobacco leaves were baked and divided into three parts, including upper leaves (the 1st to 6th leaves), middle leaves (the 7th to 12th leaves), and lower leaves (the 13th to 18th leaves). Physical properties including leaf length (cm), leaf width (cm), leaf area (cm&#xb2;), single leaf weight (g), density of leaf(g/m<sup>2</sup>), leaf moisture content (%), leaf midrib proportion (%), thickness (mm) and opening degree (%) were measured. After physical property assessment, economic traits such as yields, output value, and average price, proportion of high-quality tobacco leaves and proportion of medium to high-quality tobacco leaves were evaluated. Then, the tobacco samples were ground into fine powders for chemical composition analysis, including total sugar, reducing sugar, total N, nicotine, K, chlorine, starch, and protein. At the same time, 10 plants with consistent growth and no pests or diseases were randomly selected from each plot. The plants were dug out, cleaned, and divided into five parts: roots, stems, upper leaves, middle leaves, and lower leaves. Biomass and nutrient content for each part were subsequently measured.</p>
</sec>
<sec id="s2_3">
<label>2.3</label>
<title>Measurement methods</title>
<sec id="s2_3_1">
<label>2.3.1</label>
<title>Soil</title>
<p>The soil samples were measured using the method as described by <xref ref-type="bibr" rid="B33">Schaller et&#xa0;al. (2021)</xref> and <xref ref-type="bibr" rid="B17">Islam et&#xa0;al. (2023)</xref>. In brief, the soil pH was determined using a pH meter (ORION3STAR, Thermo Fisher Scientific, USA) in a 1:25 (w/v) paste with deionized water. OM and AN were quantified using high-temperature potassium dichromate oxidation and the alkaline hydrolysis diffusion method, respectively. AP was extracted using the Olsen method, and readily AK was extracted using the ammonium acetate extraction-flame photometry method. TN was extracted using the Kjeldahl nitrogen determination method, TP was extracted using the acid dissolution molybdenum antimony resistance colorimetric method, TK was extracted using acid dissolution flame photometry method. Trace elements in soil and plant specimens were analyzed using inductively coupled plasma mass spectrometry (ICP-MS, 7800, Agilent Technologies), Cl<sup>-</sup> using ion chromatography (IC) method (<xref ref-type="bibr" rid="B26">Mishra et&#xa0;al., 2018</xref>).</p>
</sec>
<sec id="s2_3_2">
<label>2.3.2</label>
<title>Agronomic traits</title>
<p>The agronomic traits of tobacco were measured according to the Chinese tobacco industry standard YC/T 142-1988.</p>
<p>The leaf area is calculated using the following formula:</p>
<disp-formula>
<mml:math display="block" id="M1">
<mml:mrow>
<mml:mtext>leaf&#xa0;area</mml:mtext>
<mml:mo stretchy="false">(</mml:mo>
<mml:msup>
<mml:mrow>
<mml:mtext>cm</mml:mtext>
</mml:mrow>
<mml:mn>2</mml:mn>
</mml:msup>
<mml:mo stretchy="false">)</mml:mo>
<mml:mo>=</mml:mo>
<mml:mtext>leaf&#xa0;length</mml:mtext>
<mml:mo>&#xd7;</mml:mo>
<mml:mtext>leaf&#xa0;width</mml:mtext>
<mml:mo>&#xd7;</mml:mo>
<mml:mn>0.6345.</mml:mn>
</mml:mrow>
</mml:math>
</disp-formula>
</sec>
<sec id="s2_3_3">
<label>2.3.3</label>
<title>Biomass and nutrient content</title>
<p>Tobacco samples was dried at 105&#xb0;C for half an hour and then dried at 70&#xb0;C to constant weight to determine the biomass. Roots, stems, and upper leaves, middle leaves, lower leaves were made into analytical samples by grinding with a plant crusher, and then processing through a 0.5&#xa0;mm plastic sieve.</p>
<p>The N content was determined using a Kjeldahl N analyze (<xref ref-type="bibr" rid="B37">Thilakarathna et&#xa0;al., 2019</xref>), the P content was analyzed via the molybdenum-antimony colorimetric method (<xref ref-type="bibr" rid="B44">Zhang et&#xa0;al., 2020</xref>), and the K content was measured with an FP640 flame photometer (<xref ref-type="bibr" rid="B3">Artyszak and Gozdowski, 2021</xref>).</p>
<p>The nutrient accumulation and fertilizer utilization rate are calculated using the following formula:</p>
<disp-formula>
<mml:math display="block" id="M2">
<mml:mrow>
<mml:mtext>Nutrient&#xa0;accumulation</mml:mtext>
<mml:mo stretchy="false">(</mml:mo>
<mml:mtext>g</mml:mtext>
<mml:mo stretchy="false">/</mml:mo>
<mml:mtext>plant</mml:mtext>
<mml:mo stretchy="false">)</mml:mo>
<mml:mo>=</mml:mo>
<mml:mo>&#x2211;</mml:mo>
<mml:mo stretchy="false">(</mml:mo>
<mml:mtext>nutrient&#xa0;content&#xa0;of&#xa0;each&#xa0;part</mml:mtext>
<mml:mo>&#xd7;</mml:mo>
<mml:mtext>biomass&#xa0;of&#xa0;each&#xa0;part</mml:mtext>
<mml:mo stretchy="false">)</mml:mo>
<mml:mo>;</mml:mo>
</mml:mrow>
</mml:math>
</disp-formula>
<disp-formula>
<mml:math display="block" id="M3">
<mml:mrow>
<mml:mtable equalrows="true" equalcolumns="true">
<mml:mtr>
<mml:mtd>
<mml:mrow>
<mml:mtext>Fertilizer&#xa0;utilization&#xa0;rate</mml:mtext>
<mml:mo stretchy="false">(</mml:mo>
<mml:mo>%</mml:mo>
<mml:mo stretchy="false">)</mml:mo>
<mml:mo>=</mml:mo>
<mml:mo stretchy="false">(</mml:mo>
<mml:mtext>amount&#xa0;of&#xa0;nutrients&#xa0;absorbed&#xa0;by&#xa0;crops&#xa0;in&#xa0;fertilization&#xa0;area</mml:mtext>
<mml:mo>&#x2212;</mml:mo>
</mml:mrow>
</mml:mtd>
</mml:mtr>
<mml:mtr>
<mml:mtd>
<mml:mrow>
<mml:mtext>amount&#xa0;of&#xa0;nutrients&#xa0;absorbed&#xa0;by&#xa0;crops&#xa0;in&#xa0;nutrient&#xa0;deficient&#xa0;area</mml:mtext>
<mml:mo stretchy="false">)</mml:mo>
<mml:mo stretchy="false">/</mml:mo>
<mml:mtext>amount&#xa0;of&#xa0;nutrients&#xa0;used</mml:mtext>
<mml:mo>&#xd7;</mml:mo>
<mml:mn>100</mml:mn>
<mml:mo>%</mml:mo>
</mml:mrow>
</mml:mtd>
</mml:mtr>
</mml:mtable>
</mml:mrow>
</mml:math>
</disp-formula>
</sec>
<sec id="s2_3_4">
<label>2.3.4</label>
<title>SPAD value</title>
<p>The SPAD value of tobacco plants were measured by using the Spad-520 PLUS portable chlorophyll meter.</p>
</sec>
<sec id="s2_3_5">
<label>2.3.5</label>
<title>Economic character</title>
<p>The economic characters of tobacco were calculated according to the national standard (GB2635-92).</p>
</sec>
<sec id="s2_3_6">
<label>2.3.6</label>
<title>Physical properties and chemical composition</title>
<p>The physical properties and chemical composition were measured according to the Chinese tobacco industry standard YC/T 174-2010.</p>
<p>The leaf moisture proportion, opening degree, leaf midrib proportion and density of leaves are calculated using the following formula:</p>
<disp-formula>
<mml:math display="block" id="M4">
<mml:mrow>
<mml:mtext>The&#xa0;leaf&#xa0;moisture&#xa0;proportion</mml:mtext>
<mml:mo stretchy="false">(</mml:mo>
<mml:mo>%</mml:mo>
<mml:mo stretchy="false">)</mml:mo>
<mml:mo>=</mml:mo>
<mml:mo stretchy="false">(</mml:mo>
<mml:mtext>fresh&#xa0;sample&#xa0;mass</mml:mtext>
<mml:mo>&#x2212;</mml:mo>
<mml:mtext>dry&#xa0;sample&#xa0;mass</mml:mtext>
<mml:mo stretchy="false">)</mml:mo>
<mml:mo stretchy="false">/</mml:mo>
<mml:mtext>fresh&#xa0;sample&#xa0;mass</mml:mtext>
<mml:mo>&#xd7;</mml:mo>
<mml:mn>100</mml:mn>
<mml:mo>%</mml:mo>
</mml:mrow>
</mml:math>
</disp-formula>
<disp-formula>
<mml:math display="block" id="M5">
<mml:mrow>
<mml:mtext>The&#xa0;opening&#xa0;degree</mml:mtext>
<mml:mo stretchy="false">(</mml:mo>
<mml:mo>%</mml:mo>
<mml:mo stretchy="false">)</mml:mo>
<mml:mo>=</mml:mo>
<mml:mo stretchy="false">(</mml:mo>
<mml:mtext>leaf&#xa0;width</mml:mtext>
<mml:mo stretchy="false">/</mml:mo>
<mml:mtext>leaf&#xa0;length</mml:mtext>
<mml:mo stretchy="false">)</mml:mo>
<mml:mo>&#xd7;</mml:mo>
<mml:mn>100</mml:mn>
<mml:mo>%</mml:mo>
<mml:mo>,</mml:mo>
</mml:mrow>
</mml:math>
</disp-formula>
<disp-formula>
<mml:math display="block" id="M6">
<mml:mrow>
<mml:mtext>The&#xa0;leaf&#xa0;midrib&#xa0;proportion</mml:mtext>
<mml:mo stretchy="false">(</mml:mo>
<mml:mo>%</mml:mo>
<mml:mo stretchy="false">)</mml:mo>
<mml:mo>=</mml:mo>
<mml:mo stretchy="false">(</mml:mo>
<mml:mtext>total&#xa0;mass&#xa0;of&#xa0;</mml:mtext>
<mml:mn>10</mml:mn>
<mml:mtext>&#xa0;main&#xa0;vein&#xa0;stems</mml:mtext>
<mml:mo stretchy="false">/</mml:mo>
<mml:mtext>total&#xa0;mass&#xa0;of&#xa0;</mml:mtext>
<mml:mn>10</mml:mn>
<mml:mtext>&#xa0;tobacco&#xa0;leaves</mml:mtext>
<mml:mo stretchy="false">)</mml:mo>
<mml:mo>&#xd7;</mml:mo>
<mml:mn>100</mml:mn>
</mml:mrow>
</mml:math>
</disp-formula>
<disp-formula>
<mml:math display="block" id="M7">
<mml:mrow>
<mml:mtext>The&#xa0;density&#xa0;of&#xa0;leaf</mml:mtext>
<mml:mo stretchy="false">(</mml:mo>
<mml:mtext>g</mml:mtext>
<mml:mo stretchy="false">/</mml:mo>
<mml:msup>
<mml:mrow>
<mml:mtext>m</mml:mtext>
</mml:mrow>
<mml:mn>2</mml:mn>
</mml:msup>
<mml:mo stretchy="false">)</mml:mo>
<mml:mo>=</mml:mo>
<mml:mtext>sample&#xa0;weight</mml:mtext>
<mml:mo stretchy="false">/</mml:mo>
<mml:mo stretchy="false">(</mml:mo>
<mml:mtext>number&#xa0;of&#xa0;leaf&#xa0;pieces</mml:mtext>
<mml:mo>&#xd7;</mml:mo>
<mml:mtext>single&#xa0;piece&#xa0;area</mml:mtext>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:math>
</disp-formula>
<p>Chemical compositions including total sugar, reducing sugar, total N, nicotine, water-soluble chlorine, starch, and protein were analyzed using an AA3 continuous flow analyzer, while K content was determined by flame spectrophotometry (<xref ref-type="bibr" rid="B31">Ruan et&#xa0;al., 2021</xref>).</p>
</sec>
</sec>
<sec id="s2_4">
<label>2.4</label>
<title>Statistical analysis</title>
<p>Microsoft Excel 2019 software was used to sort out data, and they were statistically analyzed using analysis of variance (ANOVA) in SPSS version 22.0 software (IBM, Chicago, USA). The significant difference among treatments was calculated by the least significant difference (LSD) and was considered significant when p&lt; 0.05.&#x2003;</p>
</sec>
</sec>
<sec id="s3" sec-type="results">
<label>3</label>
<title>Results</title>
<sec id="s3_1">
<label>3.1</label>
<title>The effect of silicon on tobacco growth and SPAD value</title>
<p>To investigate the impact of different Si fertilizer application rates on the growth of tobacco at different stages, agronomic traits including plant height, stem circumference, effective leaf number, maximum leaf length, and maximum leaf width were measured during the flower bud stage and button stage of tobacco. Results showed that, compared with S0, the plant height of S50, S100, and S200 during the flower bud stage of tobacco significantly increased by 6.05%, 16.57%, and 9.74%, respectively (<xref ref-type="table" rid="T2">
<bold>Table&#xa0;2</bold>
</xref>). Similarly, during the button stage, this phenomenon also occurred. Compared with S0, the tobacco plant height significantly increased under Si fertilizer treatment, with increases of 15.38%, 16.35%, and 19%, respectively (<xref ref-type="table" rid="T2">
<bold>Table&#xa0;2</bold>
</xref>). This indicated that Si fertilizer significantly increased the height of tobacco plants. It is worth noting that for tobacco leaves, under Si fertilizer treatment, the maximum leaf length and maximum leaf area of tobacco in the flower bud stage were significantly increased; interestingly, the maximum leaf length of tobacco during the button stage significantly increased by 7.78%, 9.6%, and 6.03%, respectively, while the maximum leaf area was showed the same trend (<xref ref-type="table" rid="T2">
<bold>Table&#xa0;2</bold>
</xref>). Compared with S0, the maximum leaf width of S200 has significantly increased, with an increase of 7.12% and 15.57% in the flower bud stage and button stage, respectively (<xref ref-type="table" rid="T2">
<bold>Table&#xa0;2</bold>
</xref>). This indicated that the application of Si fertilizer significantly promoted the growth of tobacco leaves. The effective number of tobacco leaves were increased, with significant increases of 10.20% and 13.33% in S50 and S100 during the bud stage, respectively (<xref ref-type="table" rid="T2">
<bold>Table&#xa0;2</bold>
</xref>). However, Si fertilizer had no significant effect on the stem circumference of tobacco. In summary, Si fertilizer promotes the growth and development of tobacco leaves and plants.</p>
<table-wrap id="T2" position="float">
<label>Table&#xa0;2</label>
<caption>
<p>The effects of different application rates of silicon fertilizer on the agronomic traits of tobacco.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="center">Treatments</th>
<th valign="middle" align="center">Time</th>
<th valign="middle" align="center">Plant height (cm)</th>
<th valign="middle" align="center">Stem circumference (cm)</th>
<th valign="middle" align="center">Effective number of leaves (leaves/plant)</th>
<th valign="middle" align="center">Maximum leaf length (cm)</th>
<th valign="middle" align="center">Maximum leaf width (cm)</th>
<th valign="middle" align="center">Maximum leaf area (cm<sup>2</sup>)</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S0</td>
<td valign="middle" rowspan="4" align="center" style="background-color:#ffffff">flower budding stage</td>
<td valign="middle" align="center">84.32 &#xb1; 0.45d</td>
<td valign="middle" align="center">6.44 &#xb1; 0.34a</td>
<td valign="middle" align="center">15.00 &#xb1; 0.67b</td>
<td valign="middle" align="center">50.73 &#xb1; 0.07c</td>
<td valign="middle" align="center">24.44 &#xb1; 0.44b</td>
<td valign="middle" align="center">786.87 &#xb1; 14.29b</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S50</td>
<td valign="middle" align="center">89.42 &#xb1; 1.95c</td>
<td valign="middle" align="center">6.77 &#xb1; 0.27a</td>
<td valign="middle" align="center">16.53 &#xb1; 0.24a</td>
<td valign="middle" align="center">56.09 &#xb1; 0.96b</td>
<td valign="middle" align="center">24.99 &#xb1; 0.47b</td>
<td valign="middle" align="center">889.30 &#xb1; 31.07a</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S100</td>
<td valign="middle" align="center">98.29 &#xb1; 1.14a</td>
<td valign="middle" align="center">6.84 &#xb1; 0.20a</td>
<td valign="middle" align="center">17.00 &#xb1; 0.33a</td>
<td valign="middle" align="center">58.74 &#xb1; 0.51a</td>
<td valign="middle" align="center">24.50 &#xb1; 0.58b</td>
<td valign="middle" align="center">913.04 &#xb1; 15.91a</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S200</td>
<td valign="middle" align="center">92.53 &#xb1; 1.16b</td>
<td valign="middle" align="center">6.63 &#xb1; 0.29a</td>
<td valign="middle" align="center">15.22 &#xb1; 0.19b</td>
<td valign="middle" align="center">55.25 &#xb1; 1.22b</td>
<td valign="middle" align="center">26.18 &#xb1; 0.66a</td>
<td valign="middle" align="center">917.79 &#xb1; 33.46a</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S0</td>
<td valign="middle" rowspan="4" align="center" style="background-color:#ffffff">button stage</td>
<td valign="middle" align="center">89.96 &#xb1; 2.91b</td>
<td valign="middle" align="center">6.79 &#xb1; 0.21a</td>
<td valign="middle" align="center">16.67 &#xb1; 1.00a</td>
<td valign="middle" align="center">53.47 &#xb1; 1.46b</td>
<td valign="middle" align="center">22.22 &#xb1; 1.01b</td>
<td valign="middle" align="center">748.60 &#xb1; 43.81b</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S50</td>
<td valign="middle" align="center">103.80 &#xb1; 3.90a</td>
<td valign="middle" align="center">7.08 &#xb1; 0.39a</td>
<td valign="middle" align="center">17.67 &#xb1; 1.00a</td>
<td valign="middle" align="center">57.92 &#xb1; 1.20a</td>
<td valign="middle" align="center">20.30 &#xb1; 0.52b</td>
<td valign="middle" align="center">756.24 &#xb1; 25.91b</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S100</td>
<td valign="middle" align="center">104.67 &#xb1; 2.69a</td>
<td valign="middle" align="center">6.87 &#xb1; 0.09a</td>
<td valign="middle" align="center">17.00 &#xb1; 0.33a</td>
<td valign="middle" align="center">58.90 &#xb1; 1.31a</td>
<td valign="middle" align="center">21.04 &#xb1; 1.13b</td>
<td valign="middle" align="center">786.62 &#xb1; 50.09b</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S200</td>
<td valign="middle" align="center">107.05 &#xb1; 8.59a</td>
<td valign="middle" align="center">7.03 &#xb1; 0.25a</td>
<td valign="middle" align="center">17.61 &#xb1; 0.35a</td>
<td valign="middle" align="center">56.98 &#xb1; 2.08a</td>
<td valign="middle" align="center">25.68 &#xb1; 1.92a</td>
<td valign="middle" align="center">888.32 &#xb1; 64.52a</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>The data are the mean &#xb1; standard deviation of three replicates, and different letters after values in the same column indicate significant difference by ANOVA test (P &lt; 0.05).</p>
</fn>
</table-wrap-foot>
</table-wrap>
<p>The SPAD value of tobacco leaves was measured during the bud stage and dome stage. The results indicated that, at the flower bud stage, compared with S0, Si improved the SPAD value of tobacco, and the SPAD value of tobacco leaves in S100 was significantly increased. At the button stage of tobacco, there was no significant difference among the treatments (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3</bold>
</xref>). Collectively, Si was significantly impacted the SPAD value of tobacco in the early stage.</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>SPAD values in the different treatments. S0, conventional fertilization; S50, apply 750&#xa0;kg of silicon fertilizer per hectare; S100, apply 1500&#xa0;kg of silicon fertilizer per hectare; S200, apply 3000&#xa0;kg of silicon fertilizer per hectare. The analysis of standard error and significant difference in the figure is based on the same variety. Data are means &#xb1; SE (n=4). During specific stage of tobacco, the significant difference among treatments is shown by different small letters on the error bars according to the least significant difference test (p&lt; 0.05).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1641798-g003.tif">
<alt-text content-type="machine-generated">Bar chart comparing SPAD values for Flower Bud Stage and Button Stage across treatments S0, S50, S100, and S200. Flower Bud Stage bars show values around 45 with a significant difference noted by letters a and b. Button Stage bars are slightly higher, around 50, all marked with letter a, indicating no significant difference.</alt-text>
</graphic>
</fig>
</sec>
<sec id="s3_2">
<label>3.2</label>
<title>The effect of silicon on tobacco biomass</title>
<p>During maturating stage of tobacco, the biomass of different parts of tobacco were measured, it indicated that Si fertilizer was significantly increasing the biomass of various parts of tobacco. Compared with the control, Si fertilizer significantly increased the biomass of tobacco plants, with S50, S100, and S200 significantly increasing by 26.53%, 19.95% and 19.50%, respectively (<xref ref-type="table" rid="T3">
<bold>Table&#xa0;3</bold>
</xref>). Among the upper leaves, lower leaves, stems, and roots of S50 showed the highest growth, increasing by 17.74%, 20.35%, 28.31% and 48.91%, respectively (<xref ref-type="table" rid="T3">
<bold>Table&#xa0;3</bold>
</xref>). It is interesting that the middle leaves showed the most significant increasing in S200, reaching 55.32g (<xref ref-type="table" rid="T3">
<bold>Table&#xa0;3</bold>
</xref>). Overall, these results indicated that Si fertilizer significantly increased the biomass of various parts of tobacco.</p>
<table-wrap id="T3" position="float">
<label>Table&#xa0;3</label>
<caption>
<p>The effects of different application rates of silicon fertilizer on tobacco biomass.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" rowspan="2" align="center">Treatments</th>
<th valign="middle" rowspan="2" align="center">Time</th>
<th valign="middle" colspan="6" align="center">Biomass of tobacco plants (g)</th>
<th valign="middle" colspan="6" align="center">Proportion of plant biomass growth (%)</th>
</tr>
<tr>
<th valign="middle" align="center">The whole plant</th>
<th valign="middle" align="center">Upper leaves</th>
<th valign="middle" align="center">Middle leaves</th>
<th valign="middle" align="center">Lower leaves</th>
<th valign="middle" align="center">Stem</th>
<th valign="middle" align="center">Root</th>
<th valign="middle" align="center">The whole plant</th>
<th valign="middle" align="center">Upper leaves</th>
<th valign="middle" align="center">Middle leaves</th>
<th valign="middle" align="center">Lower leaves</th>
<th valign="middle" align="center">Stem</th>
<th valign="middle" align="center">Root</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="center">S0</td>
<td valign="middle" rowspan="4" align="center">maturating stage of tobacco</td>
<td valign="middle" align="center">218.80 &#xb1; 4.84b</td>
<td valign="middle" align="center">41.26 &#xb1; 1.22c</td>
<td valign="middle" align="center">43.11 &#xb1; 0.43c</td>
<td valign="middle" align="center">38.68 &#xb1; 2.74b</td>
<td valign="middle" align="center">39.38 &#xb1; 5.08b</td>
<td valign="middle" align="center">56.37 &#xb1; 6.92b</td>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">0</td>
</tr>
<tr>
<td valign="middle" align="center">S50</td>
<td valign="middle" align="center">276.84 &#xb1; 4.83a</td>
<td valign="middle" align="center">48.58 &#xb1; 1.51a</td>
<td valign="middle" align="center">47.24 &#xb1; 1.08b</td>
<td valign="middle" align="center">46.55 &#xb1; 1.41a</td>
<td valign="middle" align="center">50.53 &#xb1; 6.55a</td>
<td valign="middle" align="center">83.93 &#xb1; 3.91a</td>
<td valign="top" align="center">26.53</td>
<td valign="top" align="center">17.74</td>
<td valign="top" align="center">9.57</td>
<td valign="top" align="center">20.35</td>
<td valign="top" align="center">28.31</td>
<td valign="top" align="center">48.91</td>
</tr>
<tr>
<td valign="middle" align="center">S100</td>
<td valign="middle" align="center">262.45 &#xb1; 5.47a</td>
<td valign="middle" align="center">45.90 &#xb1; 0.68b</td>
<td valign="middle" align="center">47.83 &#xb1; 1.52b</td>
<td valign="middle" align="center">45.71 &#xb1; 1.82a</td>
<td valign="middle" align="center">45.27 &#xb1; 5.05ab</td>
<td valign="middle" align="center">77.75 &#xb1; 4.95a</td>
<td valign="top" align="center">19.95</td>
<td valign="top" align="center">11.25</td>
<td valign="top" align="center">10.94</td>
<td valign="top" align="center">18.17</td>
<td valign="top" align="center">14.94</td>
<td valign="top" align="center">37.94</td>
</tr>
<tr>
<td valign="middle" align="center">S200</td>
<td valign="middle" align="center">261.48 &#xb1; 17.85a</td>
<td valign="middle" align="center">45.88 &#xb1; 0.67b</td>
<td valign="middle" align="center">55.32 &#xb1; 2.77a</td>
<td valign="middle" align="center">42.78 &#xb1; 2.00a</td>
<td valign="middle" align="center">48.62 &#xb1; 2.40ab</td>
<td valign="middle" align="center">68.88 &#xb1; 15.63ab</td>
<td valign="top" align="center">19.5</td>
<td valign="top" align="center">11.21</td>
<td valign="top" align="center">28.3</td>
<td valign="top" align="center">10.59</td>
<td valign="top" align="center">23.44</td>
<td valign="top" align="center">22.21</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>The data are the mean &#xb1; standard deviation of three replicates, and different letters after values in the same column indicate significant difference by ANOVA test (P &lt; 0.05).</p>
</fn>
</table-wrap-foot>
</table-wrap>
</sec>
<sec id="s3_3">
<label>3.3</label>
<title>The effect of silicon on tobacco nutrient accumulation and fertilizer utilization efficiency</title>
<p>In order to investigate the effect of different Si fertilizer application rates on the nutrient content of tobacco, the accumulation of N, P, and K in various parts of tobacco was measured. These results indicated that compared with S0, the N accumulation in S50, S100 and S200 were increased by 31.14%, 22.16%, and 14.66%, respectively (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4A</bold>
</xref>). However, among Si treatment, the N accumulation of the lower leaves, stems, and roots in S50 showed the most significantly improvement, which were increased by 27.30%, 142.48%, and 50.15%, respectively (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4A</bold>
</xref>). This data indicated that Si was significantly influence the nitrogen content of tobacco, and the application rate of 750kg/ha Si fertilizer has the most significant effect on nitrogen accumulation in tobacco.</p>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Nutrient accumulation and use efficiency in different treatments. <bold>(A)</bold> N accumulation; <bold>(B)</bold> P accumulation; <bold>(C)</bold> K accumulation; <bold>(D)</bold> N use efficiency; <bold>(E)</bold> P use efficiency; <bold>(F)</bold> K use efficiency. CK, blank, no fertilizer; S0, conventional fertilization; S50, apply 750&#xa0;kg of silicon fertilizer per hectare; S100, apply 1500&#xa0;kg of silicon fertilizer per hectare; S200, apply 3000&#xa0;kg of silicon fertilizer per hectare. The analysis of standard error and significant difference in the figure is based on the same variety. Data are means &#xb1; SE (n=4). After specific days of transplantation, the different lowercase letters inside the stacked graph and above the error bar represent significant differences among treatments according to the least significant difference test (LSD; p&lt; 0.05).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1641798-g004.tif">
<alt-text content-type="machine-generated">Six bar charts show nutrient accumulation and use efficiency of plants across different treatments (CK, S0, S50, S100, S200). Charts (A), (B), and (C) display nitrogen, phosphorus, and potassium accumulation per plant, respectively, broken down by plant parts (root, stem, leaves). Charts (D), (E), and (F) show nitrogen, phosphorus, and potassium use efficiency percentages. Labels indicate statistical significance with letters.</alt-text>
</graphic>
</fig>
<p>Research on tobacco P content found that compared with S0, there was no significant differences in P accumulation of plant and lower leaves in S50, S100, and S200 (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4B</bold>
</xref>). The P accumulation of roots in S50, S100 and S200 were increased by 56.04%, 38.98%, and 29.14%, respectively (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4B</bold>
</xref>). In contrast, the P accumulation of upper leaves of S200 were significantly decreased by 56.72%, while the P accumulation of stem of S200 were significantly increased by 32.13%; and on middle leaves, S50 was significantly decreased by22.66% (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4B</bold>
</xref>). These results indicated that Si fertilizer significantly changed P accumulation content of tobacco.</p>
<p>The application of Si significantly increased the K accumulation in various parts of the tobacco. Compared with S0, the K accumulation in S50, S100 and S200 were significantly increased by 28.08%, 28.30%, and 25.51%, respectively (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4C</bold>
</xref>). Interestingly, the K accumulation of upper leaves in S50, S100 and S200 were no significant difference, the K accumulation of middle leaves in S50, S100 and S200 were significantly increased by 18.86%, 36.18%, and 59.54%, respectively; similarly, the K accumulation of lower leaves and stem were significantly increased (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4C</bold>
</xref>). The K accumulation of roots in S50, S100 were significantly increased by 55.40%, 50.32%, and 20.83%, respectively, reaching0.59 g/plant and 0.58 g/plant (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4C</bold>
</xref>). In summary, the nutrient content of plants was significantly affected by Si fertilizer, and the application rate of moderate Si fertilizer (750&#x2013;1500 kg/ha) is better.</p>
<p>To explore the impact on the fertilize use efficiency, the utilization efficiency of N, P and K were measured, we found that Si increased the fertilizer use efficiency. Compared to S0, the N use efficiency of S50, S100 and S200 were increased by 43.71%, 31.10%, and 27.42%, respectively (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4D</bold>
</xref>). And the K use efficiency of S50, S100 and S200 were improved by 44.25%, 44.63%, and 40.25%, respectively (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4F</bold>
</xref>). While there was no significant difference among the treatments in P use efficiency (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4E</bold>
</xref>).</p>
</sec>
<sec id="s3_4">
<label>3.4</label>
<title>The effect of silicon on tobacco economic characteristics</title>
<p>In order to elucidate the impact of Si on economic characteristics, five indicators including yield, output value, average price, proportion of high-quality tobacco leaves and proportion of medium to high-quality tobacco leaves were studied. The results showed that compared with S0, the yields of S50, S100 and S200 were increased by 15.70%, 13.32%, and 17.00% respectively; similarly, the output values were increased by 30.76%, 23.14%, and 21.39% respectively (<xref ref-type="table" rid="T4">
<bold>Table&#xa0;4</bold>
</xref>). In addition, the proportion of high-quality tobacco leaves and medium to high-quality tobacco leaves were both the highest in S50, which were 80.11% and 94.75%, respectively, the average price of S50 was significantly increased by 13.08% compared to S0 (<xref ref-type="table" rid="T4">
<bold>Table&#xa0;4</bold>
</xref>). These results indicated that Si increased the yield and output value of tobacco, improved economic benefits. Furthermore, the effect of moderate Si fertilizer application is better.</p>
<table-wrap id="T4" position="float">
<label>Table&#xa0;4</label>
<caption>
<p>The effects of different application rates of silicon fertilizer on the economic characteristics of tobacco.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="center">Treatments</th>
<th valign="middle" align="center">Yield(kg/hm<sup>2</sup>)</th>
<th valign="middle" align="center">Output value(yuan/hm<sup>2</sup>)</th>
<th valign="middle" align="center">Average price(yuan/kg)</th>
<th valign="middle" align="center">Proportion of premium tobacco leaves (%)</th>
<th valign="middle" align="center">Proportion of medium and high-quality tobacco leaves (%)</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S0</td>
<td valign="middle" align="center" style="background-color:#ffffff">2048.80 &#xb1; 73.10b</td>
<td valign="middle" align="center" style="background-color:#ffffff">64227.74 &#xb1; 3216.53b</td>
<td valign="middle" align="center" style="background-color:#ffffff">32.02 &#xb1; 0.07b</td>
<td valign="middle" align="center" style="background-color:#ffffff">65.58</td>
<td valign="middle" align="center" style="background-color:#ffffff">88.96</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S50</td>
<td valign="middle" align="center" style="background-color:#ffffff">2370.49 &#xb1; 66.76a</td>
<td valign="middle" align="center" style="background-color:#ffffff">83986.71 &#xb1; 1964.59a</td>
<td valign="middle" align="center" style="background-color:#ffffff">35.45 &#xb1; 1.40a</td>
<td valign="middle" align="center" style="background-color:#ffffff">80.11</td>
<td valign="middle" align="center" style="background-color:#ffffff">94.75</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S100</td>
<td valign="middle" align="center" style="background-color:#ffffff">2321.61 &#xb1; 5.53a</td>
<td valign="middle" align="center" style="background-color:#ffffff">79089.54 &#xb1; 5009.04a</td>
<td valign="middle" align="center" style="background-color:#ffffff">34.07 &#xb1; 2.21ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">80.29</td>
<td valign="middle" align="center" style="background-color:#ffffff">93.48</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S200</td>
<td valign="middle" align="center" style="background-color:#ffffff">2397.19 &#xb1; 35.60a</td>
<td valign="middle" align="center" style="background-color:#ffffff">77963.24 &#xb1; 956.35a</td>
<td valign="middle" align="center" style="background-color:#ffffff">32.52 &#xb1; 0.09b</td>
<td valign="middle" align="center" style="background-color:#ffffff">78.01</td>
<td valign="middle" align="center" style="background-color:#ffffff">92.29</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>The data are the mean &#xb1; standard deviation of three replicates, and different letters after values in the same column indicate significant difference by ANOVA test (P &lt; 0.05).</p>
</fn>
</table-wrap-foot>
</table-wrap>
</sec>
<sec id="s3_5">
<label>3.5</label>
<title>The effect of silicon on the physical properties of tobacco leaves</title>
<p>In order to study the effect of Si on tobacco quality, the physical properties of the upper, middle, and lower leaves of tobacco were tested, including single leaf weight, density of leaf, leaf moisture proportion, leaf midrib proportion, opening degree, thickness, leaf length, leaf width and leaf area. On upper leaves, compared with S0, the single leaf weight in S50, S100 and S200 were increased by 25.17%, 11.55%, and 10.86%, respectively; and the leaf length of tobacco were significantly increased by 7.99%, 8.84%, and 6.56%, respectively; but the density of leaves was decreased by 6.08%, 9.12%, and 18.80%, respectively, S200 was significantly decreased to 108.42 g/m<sup>2</sup> (<xref ref-type="table" rid="T5">
<bold>Table&#xa0;5</bold>
</xref>). Data showed that Si promoted the growth of upper tobacco leaves, reduce tobacco density, and improved tobacco quality. Compared with S0, the leaf midrib proportion in S50 and S100 were decreased by 10.53% and 2.40%, respectively, while the leaf midrib proportion of S200 was increased by 3.95% (<xref ref-type="table" rid="T5">
<bold>Table&#xa0;5</bold>
</xref>). This indicate that high application of Si fertilizer actually increased the stem content of upper leaves, which is not conducive to improving the usability of tobacco leaves.</p>
<table-wrap id="T5" position="float">
<label>Table&#xa0;5</label>
<caption>
<p>The effects of different application rates of silicon fertilizer on the physical properties of tobacco.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="center">Treatments</th>
<th valign="middle" align="center">Different parts of tobacco leaves</th>
<th valign="middle" align="center">Single leaf weight (g)</th>
<th valign="middle" align="center">Density of leaf (g/m<sup>2</sup>)</th>
<th valign="middle" align="center">Leaf moisture proportion (%)</th>
<th valign="middle" align="center">Leaf midrib proportion (%)</th>
<th valign="middle" align="center">Opening degree (%)</th>
<th valign="middle" align="center">Thickness (mm)</th>
<th valign="middle" align="center">Leaf length (cm)</th>
<th valign="middle" align="center">Leaf width (cm)</th>
<th valign="middle" align="center">Leaf area (cm<sup>2</sup>)</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S0</td>
<td valign="middle" rowspan="4" align="center" style="background-color:#ffffff">Upper leaves</td>
<td valign="middle" align="center" style="background-color:#ffffff">5.77 &#xb1; 0.40b</td>
<td valign="middle" align="center" style="background-color:#ffffff">133.52 &#xb1; 11.77a</td>
<td valign="middle" align="center" style="background-color:#ffffff">17.90 &#xb1; 11.53a</td>
<td valign="middle" align="center" style="background-color:#ffffff">28.10 &#xb1; 1.72ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">26.23 &#xb1; 1.69a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.20 &#xb1; 0.02a</td>
<td valign="middle" align="center" style="background-color:#ffffff">46.73 &#xb1; 0.76b</td>
<td valign="middle" align="center" style="background-color:#ffffff">12.27 &#xb1; 0.99a</td>
<td valign="middle" align="center" style="background-color:#ffffff">364.04 &#xb1; 35.29a</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S50</td>
<td valign="middle" align="center" style="background-color:#ffffff">7.23 &#xb1; 0.58a</td>
<td valign="middle" align="center" style="background-color:#ffffff">125.41 &#xb1; 16.18ab</td>
<td valign="middle" align="center">11.56 &#xb1; 0.66a</td>
<td valign="middle" align="center">25.14 &#xb1; 0.93b</td>
<td valign="middle" align="center" style="background-color:#ffffff">28.07 &#xb1; 2.85a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.19 &#xb1; 0.02a</td>
<td valign="middle" align="center" style="background-color:#ffffff">50.47 &#xb1; 1.92a</td>
<td valign="middle" align="center" style="background-color:#ffffff">14.20 &#xb1; 1.91a</td>
<td valign="middle" align="center" style="background-color:#ffffff">456.17 &#xb1; 76.57a</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S100</td>
<td valign="middle" align="center" style="background-color:#ffffff">6.44 &#xb1; 0.64ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">121.35 &#xb1; 10.48ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">12.67 &#xb1; 1.53a</td>
<td valign="middle" align="center" style="background-color:#ffffff">27.43 &#xb1; 1.49ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">27.37 &#xb1; 1.64a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.18 &#xb1; 0.02a</td>
<td valign="middle" align="center" style="background-color:#ffffff">50.87 &#xb1; 1.10a</td>
<td valign="middle" align="center" style="background-color:#ffffff">13.93 &#xb1; 1.10a</td>
<td valign="middle" align="center" style="background-color:#ffffff">450.17 &#xb1; 44.48a</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S200</td>
<td valign="middle" align="center" style="background-color:#ffffff">6.40 &#xb1; 0.26ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">108.42 &#xb1; 6.87b</td>
<td valign="middle" align="center" style="background-color:#ffffff">14.54 &#xb1; 3.36a</td>
<td valign="middle" align="center" style="background-color:#ffffff">29.21 &#xb1; 1.88a</td>
<td valign="middle" align="center" style="background-color:#ffffff">26.63 &#xb1; 1.35a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.17 &#xb1; 0.01a</td>
<td valign="middle" align="center" style="background-color:#ffffff">49.80 &#xb1; 0.60a</td>
<td valign="middle" align="center" style="background-color:#ffffff">13.27 &#xb1; 0.81a</td>
<td valign="middle" align="center" style="background-color:#ffffff">419.38 &#xb1; 30.17a</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S0</td>
<td valign="middle" rowspan="4" align="center" style="background-color:#ffffff">Middle leaves</td>
<td valign="middle" align="center" style="background-color:#ffffff">6.41 &#xb1; 0.34a</td>
<td valign="middle" align="center" style="background-color:#ffffff">104.27 &#xb1; 4.54a</td>
<td valign="middle" align="center" style="background-color:#ffffff">13.72 &#xb1; 2.17a</td>
<td valign="middle" align="center" style="background-color:#ffffff">26.17 &#xb1; 1.08a</td>
<td valign="middle" align="center" style="background-color:#ffffff">32.61 &#xb1; 1.04a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.15 &#xb1; 0.00a</td>
<td valign="middle" align="center" style="background-color:#ffffff">51.33 &#xb1; 1.50a</td>
<td valign="middle" align="center" style="background-color:#ffffff">16.73 &#xb1; 0.58b</td>
<td valign="middle" align="center" style="background-color:#ffffff">545.20 &#xb1; 30.14a</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S50</td>
<td valign="middle" align="center" style="background-color:#ffffff">7.59 &#xb1; 0.92a</td>
<td valign="middle" align="center" style="background-color:#ffffff">114.46 &#xb1; 5.98a</td>
<td valign="middle" align="center" style="background-color:#ffffff">14.01 &#xb1; 2.71a</td>
<td valign="middle" align="center" style="background-color:#ffffff">20.95 &#xb1; 1.39c</td>
<td valign="middle" align="center" style="background-color:#ffffff">34.68 &#xb1; 2.79a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.16 &#xb1; 0.01a</td>
<td valign="middle" align="center" style="background-color:#ffffff">52.07 &#xb1; 2.70a</td>
<td valign="middle" align="center" style="background-color:#ffffff">18.07 &#xb1; 1.94ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">598.38 &#xb1; 90.21a</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S100</td>
<td valign="middle" align="center" style="background-color:#ffffff">6.77 &#xb1; 0.98a</td>
<td valign="middle" align="center" style="background-color:#ffffff">106.25 &#xb1; 10.34a</td>
<td valign="middle" align="center" style="background-color:#ffffff">9.14 &#xb1; 5.14a</td>
<td valign="middle" align="center" style="background-color:#ffffff">23.93 &#xb1; 1.28ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">33.62 &#xb1; 2.60a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.15 &#xb1; 0.01a</td>
<td valign="middle" align="center" style="background-color:#ffffff">52.60 &#xb1; 1.78a</td>
<td valign="middle" align="center" style="background-color:#ffffff">17.67 &#xb1; 1.10ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">589.45 &#xb1; 38.68a</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S200</td>
<td valign="middle" align="center" style="background-color:#ffffff">7.28 &#xb1; 0.21a</td>
<td valign="middle" align="center" style="background-color:#ffffff">105.02 &#xb1; 11.85a</td>
<td valign="middle" align="center" style="background-color:#ffffff">13.00 &#xb1; 0.89a</td>
<td valign="middle" align="center" style="background-color:#ffffff">23.26 &#xb1; 1.04b</td>
<td valign="middle" align="center" style="background-color:#ffffff">35.95 &#xb1; 0.69a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.15 &#xb1; 0.01a</td>
<td valign="middle" align="center" style="background-color:#ffffff">53.40 &#xb1; 0.87a</td>
<td valign="middle" align="center" style="background-color:#ffffff">19.20 &#xb1; 0.53a</td>
<td valign="middle" align="center" style="background-color:#ffffff">650.68 &#xb1; 26.74a</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S0</td>
<td valign="middle" rowspan="4" align="center" style="background-color:#ffffff">Lower leaves</td>
<td valign="middle" align="center" style="background-color:#ffffff">5.88 &#xb1; 1.11a</td>
<td valign="middle" align="center" style="background-color:#ffffff">73.51 &#xb1; 9.94a</td>
<td valign="middle" align="center" style="background-color:#ffffff">14.47 &#xb1; 1.70a</td>
<td valign="middle" align="center" style="background-color:#ffffff">27.00 &#xb1; 0.24b</td>
<td valign="middle" align="center" style="background-color:#ffffff">38.84 &#xb1; 2.33a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.11 &#xb1; 0.01a</td>
<td valign="middle" align="center" style="background-color:#ffffff">50.20 &#xb1; 2.60b</td>
<td valign="middle" align="center" style="background-color:#ffffff">19.47 &#xb1; 0.81a</td>
<td valign="middle" align="center" style="background-color:#ffffff">620.21 &#xb1; 44.26a</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S50</td>
<td valign="middle" align="center" style="background-color:#ffffff">6.53 &#xb1; 0.66a</td>
<td valign="middle" align="center" style="background-color:#ffffff">73.79 &#xb1; 4.10a</td>
<td valign="middle" align="center" style="background-color:#ffffff">15.90 &#xb1; 1.39a</td>
<td valign="middle" align="center" style="background-color:#ffffff">29.02 &#xb1; 1.36a</td>
<td valign="middle" align="center" style="background-color:#ffffff">37.21 &#xb1; 2.02a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.10 &#xb1; 0.00a</td>
<td valign="middle" align="center" style="background-color:#ffffff">54.87 &#xb1; 2.31a</td>
<td valign="middle" align="center" style="background-color:#ffffff">20.40 &#xb1; 1.06a</td>
<td valign="middle" align="center" style="background-color:#ffffff">710.52 &#xb1; 54.84a</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S100</td>
<td valign="middle" align="center" style="background-color:#ffffff">5.71 &#xb1; 0.40a</td>
<td valign="middle" align="center" style="background-color:#ffffff">65.68 &#xb1; 5.30a</td>
<td valign="middle" align="center" style="background-color:#ffffff">14.28 &#xb1; 1.19a</td>
<td valign="middle" align="center" style="background-color:#ffffff">30.38 &#xb1; 0.17a</td>
<td valign="middle" align="center" style="background-color:#ffffff">37.79 &#xb1; 1.99a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.10 &#xb1; 0.01a</td>
<td valign="middle" align="center" style="background-color:#ffffff">52.27 &#xb1; 1.62ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">19.73 &#xb1; 0.64a</td>
<td valign="middle" align="center" style="background-color:#ffffff">654.26 &#xb1; 23.92a</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S200</td>
<td valign="middle" align="center" style="background-color:#ffffff">6.49 &#xb1; 0.67a</td>
<td valign="middle" align="center" style="background-color:#ffffff">74.17 &#xb1; 4.11a</td>
<td valign="middle" align="center" style="background-color:#ffffff">15.14 &#xb1; 0.29a</td>
<td valign="middle" align="center" style="background-color:#ffffff">29.81 &#xb1; 1.54a</td>
<td valign="middle" align="center" style="background-color:#ffffff">38.58 &#xb1; 2.27a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.10 &#xb1; 0.00a</td>
<td valign="middle" align="center" style="background-color:#ffffff">53.93 &#xb1; 1.86ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">20.80 &#xb1; 1.20a</td>
<td valign="middle" align="center" style="background-color:#ffffff">712.04 &#xb1; 52.39a</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>The data are the mean &#xb1; standard deviation of three replicates, and different letters after values in the same column indicate significant difference by ANOVA test (P &lt; 0.05).</p>
</fn>
</table-wrap-foot>
</table-wrap>
<p>On middle leaves, compared with S0, the leaf midrib proportion of S50, S100 and S200 were decreased by 19.96%, 8.58%, and 11.12%, respectively; the leaf width was increased by 7.97%, 5.58%, and 14.74%, respectively, S200 was significantly increased to 19.20&#xa0;cm; similarly, the leaf area was increased by 9.75%, 8.12%, and 19.35%, respectively (<xref ref-type="table" rid="T5">
<bold>Table&#xa0;5</bold>
</xref>). This indicates that Si can promote the growth of tobacco leaves in the middle, reduce the stem content of tobacco leaves, and then increase yield.</p>
<p>The lower leaves of tobacco were impacted by Si. On lower leaves, compared with S0, the leaf midrib proportion of S50, S100 and S200 were increased by 7.48%, 12.50%, and 10.40%, respectively; the leaf length in S50, S100 and S200 were increased by 9.30%, 4.12%, and 7.44%, respectively, S50 was significantly increased to 54.87&#xa0;cm; the leaf area in S50, S100 and S200 were increased by 14.56%, 5.49%, and 14.81%, respectively (<xref ref-type="table" rid="T5">
<bold>Table&#xa0;5</bold>
</xref>). In summary, Si affected the physical properties of tobacco leaves, promoted the growth of tobacco leaves, and increased the availability of upper and middle leaves. However, high usage of Si fertilizer actually increased the leaf midrib proportion.</p>
</sec>
<sec id="s3_6">
<label>3.6</label>
<title>The effect of silicon on the chemical composition of tobacco leaves</title>
<p>Similarly, in order to investigate the effect of Si on the quality of tobacco, chemical indicators of tobacco leaf from different parts were tested. The results indicate that Si fertilizer improved the applicability of tobacco leaves. On the upper leaves, compared with S0, the content of reducing sugar in S50 was significantly increased by 15.21%, while S100 and S200 were significantly decreased by 13.92% and 21.93%, respectively; the K content of S50, S100 and S200 were significantly increased by 6.42%, 12.10%, and 11.56%, respectively; the chlorine content and the ratio of reducing sugar to total sugar of S50 were significantly increased by 20.44% and 19.09%, respectively; the nicotine content of S200 was significantly increased by 13.99%; while the total sugar, starch content and the ratio of total sugar to nicotine of S200 were significantly decreased by 14.29%, 12.05%, and 24.42%, respectively (<xref ref-type="table" rid="T6">
<bold>Table&#xa0;6</bold>
</xref>). On the middle leaves, compared with S0, the K content of S100 and S200 were significantly increased by 21.60% and 24.40%, respectively; the nicotine of S50 was significantly increased by 18.56%, the chlorine content of S100 was significantly decreased by 11.79%; the total sugar, chlorine content and the ratio of K content to chlorine content of S200 were significantly increased by 15.50%, 26.72%, and 38.35%, respectively (<xref ref-type="table" rid="T6">
<bold>Table&#xa0;6</bold>
</xref>). On the lower leaves, compared with S0, the K content of S100 was significant increased by 8.86%; the chlorine content of S50, S100 and S200 were significantly reduced by 22.25%, 18.57%, and 20.44%, respectively; the ratio of K content to chlorine content of S50, S100 and S200 were increased by 30.56%, 33.29% and 33.15% respectively; the reducing sugar content and the ratio of reducing sugar to total sugar of S50 were significantly increased by 23.96% and 15.05%, respectively; while the starch content of S50 was significantly decreased by 18.98%; the ratio of total sugar to nicotine of S100 was decreased by 7.52%; the N content of S200 was significantly decreased by 15.26%, while the starch content of S200 was significantly increased by 15.14% (<xref ref-type="table" rid="T6">
<bold>Table&#xa0;6</bold>
</xref>).</p>
<table-wrap id="T6" position="float">
<label>Table&#xa0;6</label>
<caption>
<p>The effect of different application rates of silicon fertilizer on the chemical composition of tobacco.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="center">Treatments</th>
<th valign="middle" align="center">Different parts of tobacco leaves</th>
<th valign="middle" align="center">Nitrogen (%)</th>
<th valign="middle" align="center">Nicotine (%)</th>
<th valign="middle" align="center">Total sugar (%)</th>
<th valign="middle" align="center">Reducing sugar (%)</th>
<th valign="middle" align="center">Starch (%)</th>
<th valign="middle" align="center">Potassium (%)</th>
<th valign="middle" align="center">Chlorine (%)</th>
<th valign="middle" align="center">Protein (%)</th>
<th valign="middle" align="center">Total sugar/Nicotine</th>
<th valign="middle" align="center">Potassium/Chloride</th>
<th valign="middle" align="center">Nitrogen/Nicotine</th>
<th valign="middle" align="center">Reducing sugar/Total sugar</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S0</td>
<td valign="middle" rowspan="4" align="center" style="background-color:#ffffff">Upper leaves</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.37 &#xb1; 0.10a</td>
<td valign="middle" align="center" style="background-color:#ffffff">2.81 &#xb1; 0.05b</td>
<td valign="middle" align="center" style="background-color:#ffffff">27.85 &#xb1; 0.28a</td>
<td valign="middle" align="center" style="background-color:#ffffff">18.80 &#xb1; 0.55b</td>
<td valign="middle" align="center" style="background-color:#ffffff">2.81 &#xb1; 0.22a</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.29 &#xb1; 0.06b</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.29 &#xb1; 0.02b</td>
<td valign="middle" align="center" style="background-color:#ffffff">6.51 &#xb1; 1.68a</td>
<td valign="middle" align="center" style="background-color:#ffffff">9.93 &#xb1; 0.07a</td>
<td valign="middle" align="center" style="background-color:#ffffff">4.44 &#xb1; 0.51ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.49 &#xb1; 0.04a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.68 &#xb1; 0.03b</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S50</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.20 &#xb1; 0.17a</td>
<td valign="middle" align="center" style="background-color:#ffffff">2.65 &#xb1; 0.05b</td>
<td valign="middle" align="center" style="background-color:#ffffff">26.90 &#xb1; 1.21a</td>
<td valign="middle" align="center" style="background-color:#ffffff">21.64 &#xb1; 1.97a</td>
<td valign="middle" align="center" style="background-color:#ffffff">2.63 &#xb1; 0.16ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.37 &#xb1; 0.04a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.35 &#xb1; 0.01a</td>
<td valign="middle" align="center" style="background-color:#ffffff">5.04 &#xb1; 1.08a</td>
<td valign="middle" align="center" style="background-color:#ffffff">10.17 &#xb1; 0.64a</td>
<td valign="middle" align="center" style="background-color:#ffffff">3.91 &#xb1; 0.27b</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.46 &#xb1; 0.07a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.80 &#xb1; 0.05a</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S100</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.36 &#xb1; 0.04a</td>
<td valign="middle" align="center" style="background-color:#ffffff">2.75 &#xb1; 0.15b</td>
<td valign="middle" align="center" style="background-color:#ffffff">25.85 &#xb1; 1.84ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">16.18 &#xb1; 0.41c</td>
<td valign="middle" align="center" style="background-color:#ffffff">2.88 &#xb1; 0.04a</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.45 &#xb1; 0.02a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.31 &#xb1; 0.01ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">5.45 &#xb1; 0.69a</td>
<td valign="middle" align="center" style="background-color:#ffffff">9.40 &#xb1; 0.42a</td>
<td valign="middle" align="center" style="background-color:#ffffff">4.60 &#xb1; 0.17a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.50 &#xb1; 0.01a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.63 &#xb1; 0.03b</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S200</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.39 &#xb1; 0.00a</td>
<td valign="middle" align="center" style="background-color:#ffffff">3.20 &#xb1; 0.00a</td>
<td valign="middle" align="center" style="background-color:#ffffff">24.00 &#xb1; 1.70b</td>
<td valign="middle" align="center" style="background-color:#ffffff">14.68 &#xb1; 1.22c</td>
<td valign="middle" align="center" style="background-color:#ffffff">2.47 &#xb1; 0.13b</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.44 &#xb1; 0.03a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.32 &#xb1; 0.03b</td>
<td valign="middle" align="center" style="background-color:#ffffff">4.63 &#xb1; 1.54a</td>
<td valign="middle" align="center" style="background-color:#ffffff">7.50 &#xb1; 0.52b</td>
<td valign="middle" align="center" style="background-color:#ffffff">4.53 &#xb1; 0.29ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.43 &#xb1; 0.01a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.61 &#xb1; 0.05b</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S0</td>
<td valign="middle" rowspan="4" align="center">Middle leaves</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.17 &#xb1; 0.07a</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.73 &#xb1; 0.07b</td>
<td valign="middle" align="center" style="background-color:#ffffff">27.99 &#xb1; 1.79a</td>
<td valign="middle" align="center" style="background-color:#ffffff">24.68 &#xb1; 1.43a</td>
<td valign="middle" align="center" style="background-color:#ffffff">2.62 &#xb1; 0.19b</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.45 &#xb1; 0.04b</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.27 &#xb1; 0.01b</td>
<td valign="middle" align="center" style="background-color:#ffffff">5.48 &#xb1; 0.49a</td>
<td valign="middle" align="center" style="background-color:#ffffff">16.21 &#xb1; 0.41ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">5.31 &#xb1; 0.10bc</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.68 &#xb1; 0.06a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.88 &#xb1; 0.07a</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S50</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.24 &#xb1; 0.25a</td>
<td valign="middle" align="center" style="background-color:#ffffff">2.05 &#xb1; 0.09a</td>
<td valign="middle" align="center" style="background-color:#ffffff">28.05 &#xb1; 1.71a</td>
<td valign="middle" align="center" style="background-color:#ffffff">20.99 &#xb1; 6.08a</td>
<td valign="middle" align="center" style="background-color:#ffffff">2.45 &#xb1; 0.01b</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.62 &#xb1; 0.04b</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.28 &#xb1; 0.01b</td>
<td valign="middle" align="center" style="background-color:#ffffff">6.10 &#xb1; 1.30a</td>
<td valign="middle" align="center" style="background-color:#ffffff">13.75 &#xb1; 1.41b</td>
<td valign="middle" align="center" style="background-color:#ffffff">5.85 &#xb1; 0.15b</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.60 &#xb1; 0.11a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.74 &#xb1; 0.17a</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S100</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.20 &#xb1; 0.06a</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.68 &#xb1; 0.04b</td>
<td valign="middle" align="center" style="background-color:#ffffff">29.37 &#xb1; 4.43a</td>
<td valign="middle" align="center" style="background-color:#ffffff">24.68 &#xb1; 3.91a</td>
<td valign="middle" align="center" style="background-color:#ffffff">2.62 &#xb1; 0.10b</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.77 &#xb1; 0.03a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.24 &#xb1; 0.02c</td>
<td valign="middle" align="center" style="background-color:#ffffff">6.44 &#xb1; 1.12a</td>
<td valign="middle" align="center" style="background-color:#ffffff">17.44 &#xb1; 2.59a</td>
<td valign="middle" align="center" style="background-color:#ffffff">7.35 &#xb1; 0.56a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.71 &#xb1; 0.03a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.84 &#xb1; 0.01a</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S200</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.09 &#xb1; 0.06a</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.62 &#xb1; 0.08b</td>
<td valign="middle" align="center" style="background-color:#ffffff">31.09 &#xb1; 0.64a</td>
<td valign="middle" align="center" style="background-color:#ffffff">27.28 &#xb1; 0.91a</td>
<td valign="middle" align="center" style="background-color:#ffffff">3.02 &#xb1; 0.08a</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.81 &#xb1; 0.03a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.35 &#xb1; 0.00a</td>
<td valign="middle" align="center" style="background-color:#ffffff">5.43 &#xb1; 0.40a</td>
<td valign="middle" align="center" style="background-color:#ffffff">19.26 &#xb1; 0.93a</td>
<td valign="middle" align="center" style="background-color:#ffffff">5.22 &#xb1; 0.12c</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.67 &#xb1; 0.02a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.88 &#xb1; 0.03a</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S0</td>
<td valign="middle" rowspan="4" align="center" style="background-color:#ffffff">Lower leaves</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.10 &#xb1; 0.04a</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.65 &#xb1; 0.06a</td>
<td valign="middle" align="center" style="background-color:#ffffff">30.81 &#xb1; 2.53a</td>
<td valign="middle" align="center" style="background-color:#ffffff">21.25 &#xb1; 2.66b</td>
<td valign="middle" align="center" style="background-color:#ffffff">2.93 &#xb1; 0.06b</td>
<td valign="middle" align="center" style="background-color:#ffffff">2.33 &#xb1; 0.07b</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.37 &#xb1; 0.02a</td>
<td valign="middle" align="center" style="background-color:#ffffff">4.72 &#xb1; 1.42a</td>
<td valign="middle" align="center" style="background-color:#ffffff">18.68 &#xb1; 1.11ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">6.34 &#xb1; 0.50b</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.67 &#xb1; 0.00ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.69 &#xb1; 0.03b</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S50</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.05 &#xb1; 0.02a</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.50 &#xb1; 0.02a</td>
<td valign="middle" align="center" style="background-color:#ffffff">31.42 &#xb1; 2.02a</td>
<td valign="middle" align="center" style="background-color:#ffffff">26.34 &#xb1; 3.47a</td>
<td valign="middle" align="center" style="background-color:#ffffff">2.38 &#xb1; 0.22c</td>
<td valign="middle" align="center" style="background-color:#ffffff">2.37 &#xb1; 0.11b</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.29 &#xb1; 0.03b</td>
<td valign="middle" align="center" style="background-color:#ffffff">5.31 &#xb1; 0.30a</td>
<td valign="middle" align="center" style="background-color:#ffffff">20.90 &#xb1; 1.14a</td>
<td valign="middle" align="center" style="background-color:#ffffff">8.27 &#xb1; 0.55a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.70 &#xb1; 0.01a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.84 &#xb1; 0.06a</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S100</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.06 &#xb1; 0.06a</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.66 &#xb1; 0.16a</td>
<td valign="middle" align="center" style="background-color:#ffffff">28.71 &#xb1; 3.89a</td>
<td valign="middle" align="center" style="background-color:#ffffff">18.63 &#xb1; 1.72b</td>
<td valign="middle" align="center" style="background-color:#ffffff">3.10 &#xb1; 0.09ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">2.54 &#xb1; 0.07a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.30 &#xb1; 0.01b</td>
<td valign="middle" align="center" style="background-color:#ffffff">5.17 &#xb1; 0.48a</td>
<td valign="middle" align="center" style="background-color:#ffffff">17.27 &#xb1; 1.10b</td>
<td valign="middle" align="center" style="background-color:#ffffff">8.45 &#xb1; 0.21a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.64 &#xb1; 0.08ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.65 &#xb1; 0.03b</td>
</tr>
<tr>
<td valign="middle" align="center" style="background-color:#ffffff">S200</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.93 &#xb1; 0.04b</td>
<td valign="middle" align="center" style="background-color:#ffffff">1.58 &#xb1; 0.21a</td>
<td valign="middle" align="center" style="background-color:#ffffff">32.13 &#xb1; 0.58a</td>
<td valign="middle" align="center" style="background-color:#ffffff">23.49 &#xb1; 1.61ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">3.38 &#xb1; 0.30a</td>
<td valign="middle" align="center" style="background-color:#ffffff">2.48 &#xb1; 0.05ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.29 &#xb1; 0.01b</td>
<td valign="middle" align="center" style="background-color:#ffffff">4.68 &#xb1; 0.37a</td>
<td valign="middle" align="center" style="background-color:#ffffff">20.61 &#xb1; 2.80ab</td>
<td valign="middle" align="center" style="background-color:#ffffff">8.44 &#xb1; 0.02a</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.60 &#xb1; 0.05b</td>
<td valign="middle" align="center" style="background-color:#ffffff">0.73 &#xb1; 0.04b</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>The data are the mean &#xb1; standard deviation of three replicates, and different letters after values in the same column indicate significant difference by ANOVA test (P &lt; 0.05).</p>
</fn>
</table-wrap-foot>
</table-wrap>
</sec>
</sec>
<sec id="s4" sec-type="discussion">
<label>4</label>
<title>Discussion</title>
<sec id="s4_1">
<label>4.1</label>
<title>Tobacco growth</title>
<p>Chlorophyll, as an essential pigment for photosynthesis, directly affects leaf photosynthetic capacity and organic matter accumulation (<xref ref-type="bibr" rid="B45">Zhang et&#xa0;al., 2018</xref>). Studies indicated that SPAD values were positively correlated with chlorophyll content and dry matter accumulation (<xref ref-type="bibr" rid="B6">Condorelli et&#xa0;al., 2018</xref>). <xref ref-type="bibr" rid="B25">Marxen et&#xa0;al. (2016)</xref> demonstrated that Si enhanced light interaction in plants. Researchers showed that Si application in rice and maize increased the maximum leaf area, aboveground dry matter weight and yields (<xref ref-type="bibr" rid="B29">Pati et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B5">Biswal et&#xa0;al., 2024</xref>.). In this study, the plant height and maximum leaf area of tobacco were increased under silicon treatment (<xref ref-type="table" rid="T2">
<bold>Table&#xa0;2</bold>
</xref>).</p>
<p>This study confirmed that Si fertilizer treatment significantly increased the SPAD values and biomass of tobacco (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3</bold>
</xref>; <xref ref-type="table" rid="T3">
<bold>Table&#xa0;3</bold>
</xref>). At the flower bud stage, the S100 exhibited significantly higher SPAD values than other treatments, although no significant differences were observed among the remaining treatments, their SPAD values remained numerically higher than S0 (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3</bold>
</xref>). That indicated that moderate Si supplementation improved photosynthetic capacity by elevating chlorophyll content, improved tobacco growth, the plants height and leaf area were significantly increased (<xref ref-type="bibr" rid="B16">Huang et&#xa0;al., 2024</xref>). This effect may be attributed to silicon&#x2019;s ability to optimize light utilization through increased leaf conductivity (<xref ref-type="bibr" rid="B40">Verma et&#xa0;al., 2020b</xref>). Besides, moderate Si application promoted systemic growth by stimulating root development and nutrient absorption (<xref ref-type="bibr" rid="B28">Parecido et&#xa0;al., 2025</xref>). The whole plant biomass in S50, S100 and S200 were significantly increased by 26.53%, 19.95%, and 19.50%, respectively (<xref ref-type="table" rid="T3">
<bold>Table&#xa0;3</bold>
</xref>). However, at the button stage of tobacco, there was no significant differences in SPAD values, which likely due to natural leaf senescence during later growth stages, which reduced silicon&#x2019;s chlorophyll synthesis benefits (<xref ref-type="bibr" rid="B24">Luyckx et&#xa0;al., 2017</xref>). In addition, excessive application of Si fertilizer may lead to thicker leaves, decreased stomatal conductance, affected CO<sub>2</sub> absorption and photosynthetic efficiency, decreased SPAD values, poor root development and stunted plants (<xref ref-type="bibr" rid="B16">Huang et&#xa0;al., 2024</xref>). But this phenomenon was not found in this study (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3</bold>
</xref>).</p>
</sec>
<sec id="s4_2">
<label>4.2</label>
<title>Nutrient use efficiency</title>
<sec id="s4_2_1">
<label>4.2.1</label>
<title>Nitrogen</title>
<p>It demonstrated that the application of Si enhanced N availability, promoted plant N uptake, and N use efficiency (<xref ref-type="bibr" rid="B8">da Silva et&#xa0;al., 2023</xref>; <xref ref-type="bibr" rid="B27">Neu et&#xa0;al., 2017</xref>). The results of this study were similar (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4A</bold>
</xref>). The application of Si promoted the N metabolism related microorganisms in the soil, thereby improving N availability, which was beneficial for plants to absorb N from the soil (<xref ref-type="bibr" rid="B22">Liang et&#xa0;al., 2021</xref>). Furthermore, Si induces the expression of NO3<sup>-</sup> transporter in roots, thereby improving N absorption and use efficiency, (<xref ref-type="bibr" rid="B27">Neu et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B46">Zou et al., 2018</xref>), finally, increased the accumulation of N in plants (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4A</bold>
</xref>). Similarly, in this study, the N use efficiency was significantly increased by application of Si (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4D</bold>
</xref>).</p>
</sec>
<sec id="s4_2_2">
<label>4.2.2</label>
<title>Phosphorus</title>
<p>The application of Si fertilizer at a rate of 750 kg/ha increased the P accumulation in tobacco (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4B</bold>
</xref>), similar results was observed in wheat (Triticum aestivum cv. Benchmark) and maize (<xref ref-type="bibr" rid="B13">Gunnarsen et&#xa0;al., 2022</xref>). The application of Si fertilizer reduced the soil&#x2019;s P retention capacity, leading to an increase in the level of water-soluble P thereby increasing the available P content in the soil (<xref ref-type="bibr" rid="B1">Alsaeedi et&#xa0;al., 2019</xref>). Researchers found that Si application increased the activity of functional microorganisms and important metabolic enzymes related to P cycling in the soil, thereby promoting P mineralization and improving effectiveness (<xref ref-type="bibr" rid="B22">Liang et&#xa0;al., 2021</xref>).</p>
<p>The P content of winter wheat varies with changes in Si supply (<xref ref-type="bibr" rid="B14">Hang et&#xa0;al., 2024</xref>). <xref ref-type="bibr" rid="B22">Liang et&#xa0;al. (2021)</xref>found that with the increased of Si, the activities of acid phosphatase (AcPA) in soil first increase and then decrease, when the amount of Si added is below a certain threshold, it had a positive impact on plant growth and soil nutrient utilization, however, when the threshold was exceeded, high levels of Si may lead to stress responses that negatively effects on plant growth and soil nutrient utilization. That is similar to this study, with the increase of Si fertilizer application, the P accumulation and P use efficiency were first increased and then decreased (<xref ref-type="fig" rid="f4">
<bold>Figures&#xa0;4B, E</bold>
</xref>). <xref ref-type="bibr" rid="B15">Hu et&#xa0;al. (2018)</xref> found that when Si accumulates excessively in plant, it reduces the uptake of P. In this study, the P accumulation was reduced in S100 and S200 treatments, and the P content in the middle leaves of S200 treatment was significantly lower than that S0 (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4B</bold>
</xref>). This phenomenon may be attributed to two distinct mechanisms: (1) Si reduces P uptake by downregulating the expression of the P transporter gene (<xref ref-type="bibr" rid="B15">Hu et&#xa0;al., 2018</xref>), (2) excessive Si reduces P availability in the soil (<xref ref-type="bibr" rid="B33">Schaller et&#xa0;al., 2021</xref>).</p>
</sec>
<sec id="s4_2_3">
<label>4.2.3</label>
<title>Potassium</title>
<p>Numerous studies have demonstrated that Si application significantly enhanced K use efficiency and K accumulation in plants (<xref ref-type="bibr" rid="B4">Barreto et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B28">Parecido et&#xa0;al., 2025</xref>). Consistent with these findings, our study revealed similar trends (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4</bold>
</xref>). Specifically, K use efficiency increased by 44.25% in the S50 compared to the S0 (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4F</bold>
</xref>). This improvement may be attributed to silicon&#x2019;s role in stabilizing cellular membranes and optimizing K transport channels (<xref ref-type="bibr" rid="B12">Ghosh et&#xa0;al., 2022</xref>). On the one&#xa0;hand, Si enhances K uptake by increased of aquaporin gene&#xa0;expression and aquaporin activity in plant tissues; on the other hand, the genes responsible for K transport from root cortical&#xa0;cells to the xylem were activated, thereby improving long-distance K mobilization and facilitating K translocation (<xref ref-type="bibr" rid="B4">Barreto et&#xa0;al., 2022</xref>). These synergistic mechanisms collectively explain the observed increase in K content in tobacco leaves following Si fertilizer application.</p>
</sec>
</sec>
<sec id="s4_3">
<label>4.3</label>
<title>Tobacco quality</title>
<sec id="s4_3_1">
<label>4.3.1</label>
<title>Physical properties</title>
<p>Si extends leaf growth period and promotes leaf growth by synthesizing cytokinins (<xref ref-type="bibr" rid="B10">Etesami et&#xa0;al., 2021</xref>). The study showed that the application of Si fertilizer within 750&#x2013;1500 kg/ha increased the single leaf weight of tobacco, and then, increased the yields (<xref ref-type="table" rid="T4">
<bold>Tables&#xa0;4</bold>
</xref>, <xref ref-type="table" rid="T5">
<bold>5</bold>
</xref>). However, when the application rates reached 3000 kg/ha, negative impact on single leaf weight was observed (<xref ref-type="table" rid="T5">
<bold>Table&#xa0;5</bold>
</xref>). Moderate Si application promotes cell wall thickening and leaf cell expansion, thereby increasing single-leaf weight in tobacco (<xref ref-type="bibr" rid="B40">Verma et&#xa0;al., 2020b</xref>). In contrast, excessive Si induces over-lignification of the cell wall, which restricts cellular enlargement and ultimately reduces leaf biomass (<xref ref-type="bibr" rid="B7">Coskun et&#xa0;al., 2019</xref>). Si optimizes the structure of tobacco leaves by reducing the deposition of lignin in the main veins, while excessive Si leads to an increased degree of lignification in the main veins, the leaf midrib proportion of S50 leaves was the lowest, while the leaf midrib proportion of S200 was increased (<xref ref-type="bibr" rid="B20">Kumari et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B9">El-Saadony et&#xa0;al., 2022</xref>) (<xref ref-type="table" rid="T5">
<bold>Table&#xa0;5</bold>
</xref>). In addition, excessive Si induced excessive enhancement of stem mechanical strength (<xref ref-type="bibr" rid="B33">Schaller et&#xa0;al., 2021</xref>), as evidenced by the increased leaf midrib proportion in the lower leaves of S200 (<xref ref-type="table" rid="T5">
<bold>Table&#xa0;5</bold>
</xref>).</p>
</sec>
<sec id="s4_3_2">
<label>4.3.2</label>
<title>Chemical composition</title>
<p>In this study, the effects of different amounts of Si fertilizer on different parts of tobacco leaves was vary (<xref ref-type="table" rid="T6">
<bold>Table&#xa0;6</bold>
</xref>). Si fertilization enhanced sugar conversion, improving leaf sweetness and aroma, whereas excessive application inhibited sugar-degrading enzyme activity, reducing sugar accumulation and conversion efficiency (<xref ref-type="bibr" rid="B24">Luyckx et&#xa0;al., 2017</xref>). Our results align with these findings, the upper leaves of S50 exhibited the highest reducing sugar content, correlating with optimal flavor characteristics, while the lower leaves of S200 accumulated more total sugar but displayed reduced conversion efficiency (<xref ref-type="table" rid="T6">
<bold>Table&#xa0;6</bold>
</xref>). These results responses highlight the different effects of Si application rates on leaf biochemistry.</p>
<p>
<xref ref-type="bibr" rid="B24">Luyckx et&#xa0;al. (2017)</xref> established that appropriate Si optimizes the sugar-nicotine balance, reducing leaf harshness. Our data confirm this, with S50 achieved an ideal sugar-alkaloid ratio of 10.17 in upper leaves, whereas the 3000 kg/ha application significantly decreased this ratio while increasing nicotine content (<xref ref-type="table" rid="T6">
<bold>Table&#xa0;6</bold>
</xref>), which potentially exacerbating smoking harshness.</p>
<p>Si suppressed the chloride channels (CLC family), resulted in reduced chlorine levels across Si treatments (S50, S100 and S200) relative to the control (S0) (<xref ref-type="bibr" rid="B20">Kumari et&#xa0;al., 2022</xref>). Notably, although S200 displayed marginally elevated K content, but the K-to-chlorine ratio of S100 was the more suitable value (<xref ref-type="table" rid="T6">
<bold>Table&#xa0;6</bold>
</xref>), which was attributable to excessive Si may lead to chloride ion accumulation due to soil acidification (<xref ref-type="bibr" rid="B33">Schaller et&#xa0;al., 2021</xref>). This chemical content shift coincided with decreased protein content in silicon-treated leaves (<xref ref-type="table" rid="T6">
<bold>Table&#xa0;6</bold>
</xref>), which indicating a metabolic change from N to C assimilation pathways (<xref ref-type="bibr" rid="B16">Huang et&#xa0;al., 2024</xref>). Collectively, Si fertilizer affected the quality of tobacco, and moderate Si fertilizer improved the quality of tobacco.</p>
</sec>
</sec>
<sec id="s4_4">
<label>4.4</label>
<title>Economic characteristics</title>
<p>A large number of researchers found that the application of Si increased the yields of corn, cotton, rice and sugarcane (<xref ref-type="bibr" rid="B41">Yan et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B21">Li et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B32">Sakurai et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B38">Verma et&#xa0;al., 2020a</xref>). In this study, the application of Si increased the yields of tobacco as well, which mainly due to Si promoted the growth of tobacco and increased its biomass (<xref ref-type="table" rid="T4">
<bold>Table&#xa0;4</bold>
</xref>). Additionally, moderate Si improved the physical properties and chemical balance of tobacco leaves, thus enhanced industrial usability, and the proportion of medium to high-quality tobacco was increased, besides, the proportion of medium to high-quality tobacco in the S50 treatment reached 94.75%, which increased the average price of tobacco leaves (<xref ref-type="table" rid="T4">
<bold>Table&#xa0;4</bold>
</xref>). The increase of production and average price which together lead to an increase in output value, which is beneficial for the income of tobacco farmers and the development of the industry. However, excessive investment in Si may weaken this benefit.</p>
</sec>
</sec>
<sec id="s5" sec-type="conclusions">
<label>5</label>
<title>Conclusion</title>
<p>Our study demonstrated that moderate Si application rates (750&#x2013;1500 kg/ha) significantly improved tobacco productivity and quality. Si application at 750&#x2013;1500 kg/ha increased the whole-plant biomass increased by 19.95%-26.53%, nutrient use efficiency was improved 34.83%-41.42%, high-quality leaves proportion was increased by 14.53%-14.71%, economic output value was boosted by 23.14%-30.76%. Among them, the S50 performed the best. However, excessive Si application (3000 kg/ha, S200) increased the leaf midrib proportion in the lower leaves by 10.40% and chemical imbalances was emerged, resulting in reduced of industrial availability. The study underscores the necessity of precision Si management to sustainably optimize tobacco quality, providing a scientific basis for Si fertilizer application in tobacco cultivation with practical guidance significance.</p>
</sec>
</body>
<back>
<sec id="s6" sec-type="data-availability">
<title>Data availability statement</title>
<p>The original contributions presented in the study are included in the article/supplementary material. Further inquiries can be directed to the corresponding author.</p>
</sec>
<sec id="s7" sec-type="author-contributions">
<title>Author contributions</title>
<p>MH: Conceptualization, Data curation, Software, Writing &#x2013; original draft. QL: Funding acquisition, Project administration, Resources, Writing &#x2013; review &amp; editing. YM: Formal analysis, Supervision, Writing &#x2013; review &amp; editing. PZ: Conceptualization, Writing &#x2013; review &amp; editing. KK: Formal analysis, Supervision, Writing &#x2013; original draft. BW: Conceptualization, Investigation, Writing &#x2013; review &amp; editing.</p>
</sec>
<sec id="s8" sec-type="funding-information">
<title>Funding</title>
<p>The author(s) declare that financial support was received for the research and/or publication of this article. This article obtained the funding which supplied by the project: &#x201c;Research on Key Technologies for Hunan Tobacco Quality Improvement Based on the Brand Demand of &#x2018;He Hua&#x2019; Project (HBZY2024A013)&#x201d;.</p>
</sec>
<ack>
<title>Acknowledgments</title>
<p>We are especially grateful to the reviewers and editors for appraising our manuscript.</p>
</ack>
<sec id="s9" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="s10" sec-type="ai-statement">
<title>Generative AI statement</title>
<p>The author(s) declare that no Generative AI was used in the creation of this manuscript.</p>
</sec>
<sec id="s11" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
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