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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Plant Sci.</journal-id>
<journal-title-group>
<journal-title>Frontiers in Plant Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Plant Sci.</abbrev-journal-title>
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<issn pub-type="epub">1664-462X</issn>
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<article-id pub-id-type="doi">10.3389/fpls.2026.1776194</article-id>
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<article-categories>
<subj-group subj-group-type="heading">
<subject>Original Research</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>Soil conditioners strategies for enhancing selenium bioavailability, and maize biomass in selenium-rich dryland soils</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" equal-contrib="yes">
<name><surname>Huang</surname><given-names>Taiqing</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="author-notes" rid="fn003"><sup>&#x2020;</sup></xref>
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<name><surname>Liu</surname><given-names>Shuyi</given-names></name>
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<name><surname>Huang</surname><given-names>Yanfei</given-names></name>
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<name><surname>Liu</surname><given-names>Bin</given-names></name>
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<name><surname>Tang</surname><given-names>Qizhan</given-names></name>
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<name><surname>Jiang</surname><given-names>Zepu</given-names></name>
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<aff id="aff1"><label>1</label><institution>Agricultural Resource and Environment Research Institute, Guangxi Academy of Agricultural Sciences</institution>, <city>Nanning</city>,&#xa0;<country country="cn">China</country></aff>
<aff id="aff2"><label>2</label><institution>Guangxi Academy of Agricultural Sciences</institution>, <city>Nanning</city>,&#xa0;<country country="cn">China</country></aff>
<aff id="aff3"><label>3</label><institution>Soil Fertilizer and Ecological Station of Hezhou City</institution>, <city>Hezhou</city>, <state>Guangxi</state>,&#xa0;<country country="cn">China</country></aff>
<aff id="aff4"><label>4</label><institution>Hechi Agricultural Science Research Institute</institution>, <city>Hechi</city>, <state>Guangxi</state>,&#xa0;<country country="cn">China</country></aff>
<author-notes>
<corresp id="c001"><label>*</label>Correspondence: Yanfei Huang, <email xlink:href="mailto:hyfei86@gxaas.net">hyfei86@gxaas.net</email>; Zhong Liu, <email xlink:href="mailto:liuzhong@gxaas.net">liuzhong@gxaas.net</email></corresp>
<fn fn-type="equal" id="fn003">
<label>&#x2020;</label>
<p>These authors have contributed equally to this work</p></fn>
</author-notes>
<pub-date publication-format="electronic" date-type="pub" iso-8601-date="2026-03-04">
<day>04</day>
<month>03</month>
<year>2026</year>
</pub-date>
<pub-date publication-format="electronic" date-type="collection">
<year>2026</year>
</pub-date>
<volume>17</volume>
<elocation-id>1776194</elocation-id>
<history>
<date date-type="received">
<day>27</day>
<month>12</month>
<year>2025</year>
</date>
<date date-type="accepted">
<day>02</day>
<month>02</month>
<year>2026</year>
</date>
<date date-type="rev-recd">
<day>26</day>
<month>01</month>
<year>2026</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2026 Huang, Liu, Liao, Wang, Peng, Wei, Huang, Liu, Liu, Tang and Jiang.</copyright-statement>
<copyright-year>2026</copyright-year>
<copyright-holder>Huang, Liu, Liao, Wang, Peng, Wei, Huang, Liu, Liu, Tang and Jiang</copyright-holder>
<license>
<ali:license_ref start_date="2026-03-04">https://creativecommons.org/licenses/by/4.0/</ali:license_ref>
<license-p>This is an open-access article distributed under the terms of the <ext-link ext-link-type="uri" xlink:href="https://creativecommons.org/licenses/by/4.0/">Creative Commons Attribution License (CC BY)</ext-link>. 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.</license-p>
</license>
</permissions>
<abstract>
<p>Low selenium bioavailability in selenium-rich soils represents a key constraint limiting selenium biofortification in agriculture. This study evaluated soil conditioning strategies to enhance selenium bioavailability, providing a theoretical foundation for efficient utilization of selenium-enriched soil resources. A pot experiment tested four soil conditioners: organic fertilizer, potassium humate, lime, and biochar, across three consecutive maize plantings. Soil conditioners effectively modified soil physicochemical properties: lime significantly increased pH and available phosphorus, while organic fertilizer increased available sulfur. These amendments markedly affected soluble and exchangeable selenium fractions. All treatments progressively increased the proportion of soluble selenium, with lime and biochar showing the most substantial gains in batches two and three (0.59%, 0.23%, 0.67%, and 0.30%, respectively). Organic fertilizer, lime, and biochar consistently elevated root selenium concentrations across all three batches by 7.79&#x2013;11.01%, 33.31&#x2013;135.41%, and 28.84&#x2013;40.81%, respectively. Potassium humate increased root selenium by 9.04&#x2013;26.42% in batches two and three. Notably, only lime consistently enhanced shoot selenium by 40.13&#x2013;87.38% across all batches, while biochar increased shoot selenium by 5.60% in batch three. Plant selenium translocation analysis revealed that only lime treatment in batch three significantly increased the selenium transfer coefficient. Correlation analysis demonstrated a highly significant positive relationship between shoot selenium content and soil pH, whereas root selenium showed no significant correlation with soluble or exchangeable selenium fractions. In selenium-rich dryland soils, conditioner application increases soil pH, thereby enhancing selenium availability and root absorption. Lime proved most effective for increasing crop selenium content, while biochar also substantially improved soil selenium availability.</p>
</abstract>
<kwd-group>
<kwd>maize</kwd>
<kwd>selenium content</kwd>
<kwd>soil conditioner</kwd>
<kwd>soil selenium speciation</kwd>
<kwd>transfer coefficient</kwd>
</kwd-group>
<funding-group>
<funding-statement>The author(s) declared that financial support was received for this work and/or its publication. This work was supported by the Guangxi Natural Science Foundation (2024GXNSFAA010491), National Natural Science Foundation of China (41761052), Special Program for Basic Research Operations of Guangxi Academy of Agricultural Sciences (GuiNongKe 2026YT147), and the Research and Technology Development Fund of Guangxi Academy of Agricultural Sciences (GuiNongKe 2024ZX33).</funding-statement>
</funding-group>
<counts>
<fig-count count="5"/>
<table-count count="4"/>
<equation-count count="0"/>
<ref-count count="43"/>
<page-count count="12"/>
<word-count count="6793"/>
</counts>
<custom-meta-group>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Plant Nutrition</meta-value>
</custom-meta>
</custom-meta-group>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>Selenium is an essential micronutrient for human health (<xref ref-type="bibr" rid="B2">Cao et&#xa0;al., 2025</xref>). Plant-derived selenium serves as the primary dietary source for humans (<xref ref-type="bibr" rid="B29">Rayman, 2008</xref>; <xref ref-type="bibr" rid="B11">Gu et&#xa0;al., 2025</xref>). Globally, approximately 1 billion people experience inadequate daily selenium intake (<xref ref-type="bibr" rid="B13">Haug et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B11">Gu et&#xa0;al., 2025</xref>), with this proportion reaching 36%&#x2013;61% in China (<xref ref-type="bibr" rid="B5">Dinh et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B2">Cao et&#xa0;al., 2025</xref>). Soil selenium represents the main source of plant selenium accumulation. Plant selenium content depends on the total selenium content of selenium-rich soil (<xref ref-type="bibr" rid="B38">Wu et&#xa0;al., 2025</xref>). Selenium undergoes interconversion among various chemical forms in soil through oxidation-reduction, humification, and methylation (<xref ref-type="bibr" rid="B25">Pu et&#xa0;al., 2023</xref>). Soil environmental conditions significantly influence selenium speciation, with higher available selenium content promoting selenium absorption and accumulation in crops (<xref ref-type="bibr" rid="B6">Dinh et&#xa0;al., 2019</xref>). Although total selenium content in southern China soils is generally high (<xref ref-type="bibr" rid="B43">Zhao et&#xa0;al., 2024</xref>), selenium levels in many agricultural products remain low (<xref ref-type="bibr" rid="B7">Du et&#xa0;al., 2018</xref>). Excessive selenium (Se) acts as a pro-oxidant in plants, causing significant toxicity (selenosis) characterized by stunted growth, root inhibition, chlorosis, and necrosis (<xref ref-type="bibr" rid="B11">Gu et&#xa0;al., 2025</xref>). It causes damage by replacing sulfur in amino acids (cysteine and methionine), resulting in dysfunctional proteins, alongside inducing severe oxidative stress (<xref ref-type="bibr" rid="B2">Cao et&#xa0;al., 2025</xref>). By modulating soil physical and chemical properties to improve selenium bioavailability, crop selenium absorption and accumulation can be enhanced, thereby improving human selenium nutrition status.</p>
<p>Plant roots primarily absorb water-soluble chemical selenium forms such as Se<sup>6+</sup>, Se<sup>4+</sup>, and organic selenium from soil solution (<xref ref-type="bibr" rid="B36">White, 2018</xref>). These water-soluble chemical selenium forms exist predominantly as water-soluble and partially exchangeable fractions, collectively termed extractable selenium (<xref ref-type="bibr" rid="B20">Loffredo et&#xa0;al., 2011</xref>). Studies of typical selenium-rich soils in China indicate that these two fractions account for approximately 0.7% and 5.1% of total selenium in the soil, respectively (<xref ref-type="bibr" rid="B22">Lyu et&#xa0;al., 2021</xref>). Soil pH, mineral composition, and organic matter content are closely related to soil selenium speciation characteristics (<xref ref-type="bibr" rid="B37">Winkel et&#xa0;al., 2015</xref>). Selenate (SeO<sub>4</sub><sup>2&#x2013;</sup>) is the main water-soluble form of Se in oxic soils (pH + pe &gt; 15), which include most cultivated soils, whereas selenite (SeO<sub>3</sub><sup>2&#x2013;</sup>) predominates in anaerobic soils with a neutral to acidic pH (pH + pe = 7.5&#x2013;15), such as paddy soils (<xref ref-type="bibr" rid="B9">Fordyce, 2013</xref>; <xref ref-type="bibr" rid="B24">Pilbeam et&#xa0;al., 2015</xref>). Selenide (Se<sup>2&#x2013;</sup>) species are stable only under low redox conditions (pH + pe &lt; 7.5) and are rarely present in cultivated soils. Selenate is relatively mobile in the soil solution, but selenite is strongly absorbed by iron and aluminum oxides/hydroxides and, to a lesser extent, by clays and organic matter (<xref ref-type="bibr" rid="B24">Pilbeam et&#xa0;al., 2015</xref>). Thus, the addition of selenate to soils facilitates immediate Se accumulation by plants, while selenite provides a longer lasting Se fertilizer (<xref ref-type="bibr" rid="B9">Fordyce, 2013</xref>; <xref ref-type="bibr" rid="B24">Pilbeam et&#xa0;al., 2015</xref>). Soil organic matter and iron oxides both adsorb selenium, reducing its bioavailability (<xref ref-type="bibr" rid="B26">Qin et&#xa0;al., 2023</xref>).</p>
<p>Adding soil conditioners can modify soil physicochemical properties, thereby influencing the cycling and transformation of soil elements (<xref ref-type="bibr" rid="B14">Huang et&#xa0;al., 2024</xref>). Biochar application in paddy fields reduces the selenium uptake by rice roots (<xref ref-type="bibr" rid="B15">Huang et&#xa0;al., 2019</xref>), whereas in dryland soils, it enhances plant selenium absorption (<xref ref-type="bibr" rid="B23">Ma et&#xa0;al., 2023</xref>). Organic fertilizer is often reported to decrease selenium bioavailability by adsorbing and immobilizing soil selenium (<xref ref-type="bibr" rid="B17">Li et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B30">Sharma et&#xa0;al., 2011</xref>); however, long-term field experiments show that wheat grains grown with organic fertilizer contain significantly higher selenium than those receiving chemical fertilizer (<xref ref-type="bibr" rid="B3">Daud et&#xa0;al., 2024</xref>). Returning selenium-enriched crop straw to the field can also increase selenium content in subsequent crops (<xref ref-type="bibr" rid="B33">Wang et&#xa0;al., 2018</xref>). Yet conventional straw return promotes the transformation of water-soluble small-molecule organic selenium (FA-Se) into large-molecule humic-bound selenium (HA-Se), reducing soil selenium bioavailability (<xref ref-type="bibr" rid="B34">Wang et&#xa0;al., 2020</xref>).</p>
<p>Studies using selenium-enriched soils with added exogenous selenium demonstrate that sulfur application significantly decreases water-soluble selenium while promoting its association with iron and manganese oxides and organic matter, thereby reducing plant selenium content (<xref ref-type="bibr" rid="B4">Deng et&#xa0;al., 2021</xref>). In contrast, in naturally selenium-poor soils without exogenous selenium, sulfur application can enhance wheat selenium content (<xref ref-type="bibr" rid="B40">Yeasmin et&#xa0;al., 2022</xref>). <xref ref-type="bibr" rid="B6">Dinh et&#xa0;al. (2019)</xref> showed that lime and coal slag can regulate selenium availability in acidic selenium-rich soils in southern China, but the effects of the same amendments vary by soil type, either increasing or decreasing available selenium (<xref ref-type="bibr" rid="B18">Liu et&#xa0;al., 2021</xref>). <xref ref-type="bibr" rid="B39">Xie et&#xa0;al. (2017)</xref> studied the effects of phosphorus application on soil selenium speciation and wheat plant selenium content, showing that different phosphorus sources had different effects. The application of urea phosphate and ammonium polyphosphate did not significantly alter soil selenium fractions or wheat selenium content in different organs (<xref ref-type="bibr" rid="B1">An et&#xa0;al., 2019</xref>). In contrast, lime and disodium hydrogen phosphate increased water-soluble and exchangeable selenium and decreased residual selenium, effectively improving selenium availability and uptake in rice (<xref ref-type="bibr" rid="B32">Tan et&#xa0;al., 2024</xref>).</p>
<p>Overall, the impact of soil amendments on selenium bioavailability is inconsistent, reflecting the strong influence of local soil conditions on the effectiveness of selenium-activating agricultural measures (<xref ref-type="bibr" rid="B6">Dinh et&#xa0;al., 2019</xref>). These discrepancies may arise from regional differences in the dominant factors controlling soil selenium bioavailability (<xref ref-type="bibr" rid="B8">Farooq et&#xa0;al., 2023</xref>). In southern China, soils are generally high in total selenium but low in available selenium, yet studies on selenium activation remain limited. This study focuses on typical selenium-rich red soils in this region, using maize as a representative crop, to evaluate the effects of different soil conditioners on selenium speciation transformation and bioavailability in dryland soils, providing a theoretical foundation and practical guidance for the efficient utilization of selenium-rich soils in this area.</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>Sample collection</title>
<p>The experiment was conducted in a greenhouse at the Guangxi Academy of Agricultural Sciences Research and Experiment Base from March to August 2023. Potting soil was collected from Yandong Township, Debao County, Baise City, Guangxi Province. Its basic physicochemical properties were: organic matter 30.3 g/kg, total nitrogen 1.8 g/kg, total phosphorus 1.25&#xa0;g/kg, total potassium 29.22 g/kg, available nitrogen 92 mg/kg, available phosphorus 77 mg/kg, available potassium 183 mg/kg, pH 7.0, and total selenium 0.82 mg/kg (<xref ref-type="table" rid="T1"><bold>Table&#xa0;1</bold></xref>).</p>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Soil basic physicochemical properties.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="center">pH</th>
<th valign="middle" align="center">OM (g/kg)</th>
<th valign="middle" align="center">TN (g/kg)</th>
<th valign="middle" align="center">TP (g/kg)</th>
<th valign="middle" align="center">TK (g/kg)</th>
<th valign="middle" align="center">TSe (mg/kg)</th>
<th valign="middle" align="center">AN (mg/kg)</th>
<th valign="middle" align="center">AP (mg/kg)</th>
<th valign="middle" align="center">AK (mg/kg)</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="center">7.00</td>
<td valign="middle" align="center">30.30</td>
<td valign="middle" align="center">1.80</td>
<td valign="middle" align="center">1.25</td>
<td valign="middle" align="center">29.22</td>
<td valign="middle" align="center">0.82</td>
<td valign="middle" align="center">92.00</td>
<td valign="middle" align="center">77.00</td>
<td valign="middle" align="center">183.00</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>OM, organic matter; TN, total nitrogen; TP, total phosphorus; TK, total potassium; TSe, total selenium; AN, available nitrogen; AP, available phosphorus; AK, available potassium.</p></fn>
</table-wrap-foot>
</table-wrap>
<p>The soil conditioners used were as follows: organic fertilizer containing 461 g/kg organic matter, N 29 g/kg, P<sub>2O5</sub> 11 g/kg, K<sub>2</sub>O 19&#xa0;g/kg, pH 5.5; potassium humate with 373 g/kg organic matter, N 24.1 g/kg, P<sub>205</sub> 6 g/kg, K<sub>2</sub>O 61.2 g/kg; and biochar with 423 g/kg organic C, total nitrogen 11.1 g/kg, P<sub>205</sub> 7 g/kg, K<sub>2</sub>O 32.6 g/kg, pH 9.3. The maize variety used was Guidan 162.</p>
</sec>
<sec id="s2_2">
<label>2.2</label>
<title>Experimental design</title>
<p>The experiment was performed in pots with five treatments: organic fertilizer (OF, 3.13 g/kg soil), quicklime (QL, 1.25 g/kg soil), biochar (BC, 10 g/kg soil), potassium humate (FA, 5 g/kg soil), and a conventional fertilization control (CK). Chemical fertilizers were applied at the same rate across all treatments. Each treatment was replicated four times. All soil conditioners were thoroughly mixed with the soil and then placed in the pots before planting the first batch of corn. No soil conditioners were added for the second and third batches of corn planted thereafter.</p>
<p>Biochar was thoroughly mixed with the soil before planting the first maize batch; no biochar was added for subsequent batches. Each batch received only basal fertilizer (compound fertilizer 15-15-15, 3.00&#xa0;g/pot), mixed with the soil prior to planting. Maize seeds were sown at five per pot, thinned to two plants per pot at the three-leaf stage. Irrigation was applied daily to maintain 70% of field capacity. Three consecutive maize batches were planted, each grown for 30 days.</p>
</sec>
<sec id="s2_3">
<label>2.3</label>
<title>Sample collection and analysis</title>
<sec id="s2_3_1">
<label>2.3.1</label>
<title>Sample collection and preparation</title>
<p>After each 30-day growth period, maize roots, stems, and leaves were harvested, washed, dried, weighed, and ground for total selenium analysis. Simultaneously, pot soils were thoroughly mixed and sampled for physicochemical property assessment and selenium speciation analysis.</p>
</sec>
<sec id="s2_3_2">
<label>2.3.2</label>
<title>Analytical methods</title>
<p>Soil pH was measured potentiometrically (soil-water ratio 1:2.5). Available phosphorus was determined via sodium carbonate extraction-molybdenum antimony colorimetry, and available sulfur via phosphate extraction-barium sulfate turbidimetry. Soil total selenium was quantified using hydride generation-atomic fluorescence spectrometry according to NY/T 1104-2006 (<xref ref-type="bibr" rid="B2">Cao et&#xa0;al., 2025</xref>).</p>
<p>Plant total selenium was determined following GB 5009.93-2017. Briefly, 0.3000&#xa0;g of plant material was digested in 8 mL concentrated nitric acid at 120 &#xb0;C for 30&#xa0;min, followed by microwave digestion (800 W for 10&#xa0;min, hold 5&#xa0;min; 1400 W for 10&#xa0;min, hold 20&#xa0;min) (<xref ref-type="bibr" rid="B1">An et&#xa0;al., 2019</xref>). The digest was cooled, reduced to 2 mL, treated with 5 mL 1:1 HCl, digested for 15&#xa0;min, diluted to 50 mL, and analyzed using an atomic fluorescence spectrometer (Jitian AFS-8330).</p>
<p>Soil selenium speciation was determined using a <xref ref-type="bibr" rid="B4">Deng et&#xa0;al. (2021)</xref> five-step sequential extraction method. One gram of air-dried, sieved soil (0.15&#xa0;mm) was extracted at a 1:10 soil-to-solution ratio: (i) soluble Se (Sol-Se) with 0.25 mol/L KCl at 25 &#xb0;C for 1&#xa0;h; (ii) exchangeable Se (Exc-Se) with 0.7 mol/L KH<sub>2</sub>PO<sub>4</sub> (pH 5.0) at 25 &#xb0;C for 4&#xa0;h; (iii) iron/manganese oxide-bound Se (FMO-Se) with 2.5&#xa0;mol/L HCl at 90 &#xb0;C for 4&#xa0;h; (iv) organic matter-bound Se (OM-Se) with 8 mL 5% K<sub>2</sub>S<sub>2</sub>O<sub>8</sub> + 2 mL 1:1 HNO<sub>3</sub> at 90&#xb0;C for 3&#xa0;h; (v) residual Se (Res-Se) determined on the remaining soil using NY/T 1104-2006. Supernatants were centrifuged at 4000 rpm for 10&#xa0;min and stored at 4 &#xb0;C for analysis (<xref ref-type="bibr" rid="B4">Deng et&#xa0;al., 2021</xref>).</p>
</sec>
</sec>
<sec id="s2_4">
<label>2.4</label>
<title>Data analysis</title>
<p>The data were organized using Microsoft Excel 2010 (Microsoft Corp., Raymond, Washington, USA) and SPSS version 26. ANOVA was employed to analyze the effects of differences between treatments were assessed by least significant difference (LSD) tests (<italic>P &lt;</italic> 0.05). Figures, correlation analysis and plotting were conducted using Origin 2021 (Origin Lab, Corp., Hampton, Massachusetts, USA).</p>
</sec>
</sec>
<sec id="s3" sec-type="results">
<label>3</label>
<title>Results</title>
<sec id="s3_1">
<label>3.1</label>
<title>Effects of soil conditioner application on soil pH, available phosphorus, and available sulfur</title>
<p><xref ref-type="fig" rid="f1"><bold>Figure&#xa0;1</bold></xref> illustrates the dynamics of these parameters under different soil conditioner treatments. Soil pH under lime, biochar, and conventional treatments initially increased and then gradually declined. In contrast, potassium humate treatment exhibited a decrease followed by an increase and a subsequent decline, while organic fertilizer initially reduced pH, which then gradually increased. Among all treatments, lime consistently maintained the highest pH. Except for the first maize batch under organic fertilizer, which showed significantly lower pH than biochar and conventional treatments, there were no significant pH differences among biochar, conventional, potassium humate, and organic fertilizer treatments after the second and third maize batches. Overall, pH remained higher under biochar and conventional treatments compared to potassium humate and organic fertilizer.</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Variation of soil pH, available phosphorus, and available sulfur under different soil conditioner treatments. Different lowercase letters within each batch indicate significant differences among treatments (<italic>P</italic>&#xa0;&lt;&#xa0;0.05). The vertical bars represent the standard error of the mean (n = 4).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-17-1776194-g001.tif">
<alt-text content-type="machine-generated">Three line graphs compare CK, OF, FA, QL, and BC treatments for pH values, available phosphorus, and available sulfur concentrations across three batches, with error bars and significance letters marked for each data point.</alt-text>
</graphic></fig>
<p>Available phosphorus levels were significantly higher only under the conventional treatment during the first two maize batches, while no significant differences were observed among lime, biochar, potassium humate, and organic fertilizer treatments. After the third batch, available phosphorus levels showed no significant differences among any treatments.</p>
<p>Available sulfur content was significantly higher in soils treated with organic fertilizer after the first and second maize batches, whereas other treatments showed no significant differences. In the third batch, available sulfur increased under potassium humate treatment, resulting in both organic fertilizer and potassium humate treatments having significantly higher sulfur content than the other three treatments. No significant differences were observed between organic fertilizer and potassium humate, or among lime, biochar, and conventional treatments.</p>
</sec>
<sec id="s3_2">
<label>3.2</label>
<title>Effects of soil conditioning on soil selenium speciation</title>
<p><xref ref-type="table" rid="T2"><bold>Table&#xa0;2</bold></xref> presents the proportions of soil selenium forms following application of different soil conditioners. After the first maize batch, soil conditioners primarily influenced exchangeable and residual selenium, whereas soluble selenium, iron-manganese-bound selenium, and organically bound selenium showed no significant differences among treatments. Exchangeable selenium was highest under organic fertilizer, significantly exceeding all other treatments, and lowest under the control (CK). Residual selenium was highest in the biochar treatment, surpassing lime, potassium humate, organic fertilizer, and CK by 3.91, 5.03, 5.78, and 2.97 percentage points, respectively. Available selenium proportions in organic fertilizer, potassium humate, lime, and biochar treatments were 3.79%, 2.38%, 2.56%, and 2.23%, respectively, all higher than CK.</p>
<table-wrap id="T2" position="float">
<label>Table&#xa0;2</label>
<caption>
<p>Soil selenium speciation composition under different soil conditioner treatments.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" colspan="2" align="center">Treatments</th>
<th valign="middle" align="center">Soluble-Se (%)</th>
<th valign="middle" align="center">Exchangeable-Se (%)</th>
<th valign="middle" align="center">Fe/Mn oxide-bound-Se (%)</th>
<th valign="middle" align="center">Organic matter-bond-Se (%)</th>
<th valign="middle" align="center">Residual-Se (%)</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" rowspan="5" align="center">Batch 1</td>
<td valign="middle" align="center">CK</td>
<td valign="middle" align="left">0.44&#xa0;&#xb1;&#xa0;0.04 a</td>
<td valign="middle" align="left">1.23&#xa0;&#xb1;&#xa0;0.35 c</td>
<td valign="middle" align="left">46.90&#xa0;&#xb1;&#xa0;2.95 a</td>
<td valign="middle" align="left">30.93&#xa0;&#xb1;&#xa0;0.60 a</td>
<td valign="middle" align="left">20.50&#xa0;&#xb1;&#xa0;3.18 ab</td>
</tr>
<tr>
<td valign="middle" align="center">OF</td>
<td valign="middle" align="left">0.45&#xa0;&#xb1;&#xa0;0.03 a</td>
<td valign="middle" align="left">3.34&#xa0;&#xb1;&#xa0;0.28 a</td>
<td valign="middle" align="left">45.69&#xa0;&#xb1;&#xa0;0.87 a</td>
<td valign="middle" align="left">32.83&#xa0;&#xb1;&#xa0;0.22 a</td>
<td valign="middle" align="left">17.68&#xa0;&#xb1;&#xa0;0.85 b</td>
</tr>
<tr>
<td valign="middle" align="center">FA</td>
<td valign="middle" align="left">0.52&#xa0;&#xb1;&#xa0;0.10 a</td>
<td valign="middle" align="left">1.86&#xa0;&#xb1;&#xa0;0.30 bc</td>
<td valign="middle" align="left">45.84&#xa0;&#xb1;&#xa0;1.40 a</td>
<td valign="middle" align="left">33.34&#xa0;&#xb1;&#xa0;1.16 a</td>
<td valign="middle" align="left">18.44&#xa0;&#xb1;&#xa0;1.89 b</td>
</tr>
<tr>
<td valign="middle" align="center">QL</td>
<td valign="middle" align="left">0.51&#xa0;&#xb1;&#xa0;0.07 a</td>
<td valign="middle" align="left">2.05&#xa0;&#xb1;&#xa0;0.46 b</td>
<td valign="middle" align="left">45.48&#xa0;&#xb1;&#xa0;2.38 a</td>
<td valign="middle" align="left">32.40&#xa0;&#xb1;&#xa0;2.03 a</td>
<td valign="middle" align="left">19.56&#xa0;&#xb1;&#xa0;1.91 ab</td>
</tr>
<tr>
<td valign="middle" align="center">BC</td>
<td valign="middle" align="left">0.51&#xa0;&#xb1;&#xa0;0.07 a</td>
<td valign="middle" align="left">1.72&#xa0;&#xb1;&#xa0;0.26 bc</td>
<td valign="middle" align="left">43.20&#xa0;&#xb1;&#xa0;2.09 a</td>
<td valign="middle" align="left">31.10&#xa0;&#xb1;&#xa0;0.97 a</td>
<td valign="middle" align="left">23.47&#xa0;&#xb1;&#xa0;2.27 a</td>
</tr>
<tr>
<td valign="middle" rowspan="5" align="center">Batch 2</td>
<td valign="middle" align="center">CK</td>
<td valign="middle" align="left">0.36&#xa0;&#xb1;&#xa0;0.05 b</td>
<td valign="middle" align="left">2.36&#xa0;&#xb1;&#xa0;0.23 b</td>
<td valign="middle" align="left">40.30&#xa0;&#xb1;&#xa0;4.98 a</td>
<td valign="middle" align="left">32.84&#xa0;&#xb1;&#xa0;5.88 ab</td>
<td valign="middle" align="left">24.13&#xa0;&#xb1;&#xa0;4.41 a</td>
</tr>
<tr>
<td valign="middle" align="center">OF</td>
<td valign="middle" align="left">0.36&#xa0;&#xb1;&#xa0;0.06 b</td>
<td valign="middle" align="left">0.21&#xa0;&#xb1;&#xa0;0.05 c</td>
<td valign="middle" align="left">36.28&#xa0;&#xb1;&#xa0;2.21 a</td>
<td valign="middle" align="left">38.90&#xa0;&#xb1;&#xa0;0.82 a</td>
<td valign="middle" align="left">24.26&#xa0;&#xb1;&#xa0;2.37 a</td>
</tr>
<tr>
<td valign="middle" align="center">FA</td>
<td valign="middle" align="left">0.41&#xa0;&#xb1;&#xa0;0.07 b</td>
<td valign="middle" align="left">0.38&#xa0;&#xb1;&#xa0;0.07 c</td>
<td valign="middle" align="left">38.87&#xa0;&#xb1;&#xa0;3.48 a</td>
<td valign="middle" align="left">36.56&#xa0;&#xb1;&#xa0;3.49 ab</td>
<td valign="middle" align="left">23.78&#xa0;&#xb1;&#xa0;1.48 a</td>
</tr>
<tr>
<td valign="middle" align="center">QL</td>
<td valign="middle" align="left">0.95&#xa0;&#xb1;&#xa0;0.25 a</td>
<td valign="middle" align="left">0.77&#xa0;&#xb1;&#xa0;0.41 c</td>
<td valign="middle" align="left">41.81&#xa0;&#xb1;&#xa0;1.56 a</td>
<td valign="middle" align="left">34.18&#xa0;&#xb1;&#xa0;1.56 ab</td>
<td valign="middle" align="left">22.29&#xa0;&#xb1;&#xa0;1.87 a</td>
</tr>
<tr>
<td valign="middle" align="center">BC</td>
<td valign="middle" align="left">1.03&#xa0;&#xb1;&#xa0;0.11 a</td>
<td valign="middle" align="left">5.51&#xa0;&#xb1;&#xa0;1.35 a</td>
<td valign="middle" align="left">40.05&#xa0;&#xb1;&#xa0;1.35 a</td>
<td valign="middle" align="left">30.74&#xa0;&#xb1;&#xa0;1.36 b</td>
<td valign="middle" align="left">22.68&#xa0;&#xb1;&#xa0;2.54 a</td>
</tr>
<tr>
<td valign="middle" rowspan="5" align="center">Batch 3</td>
<td valign="middle" align="center">CK</td>
<td valign="middle" align="left">0.12&#xa0;&#xb1;&#xa0;0.06 b</td>
<td valign="middle" align="left">2.76&#xa0;&#xb1;&#xa0;0.43 bc</td>
<td valign="middle" align="left">32.99&#xa0;&#xb1;&#xa0;1.84 a</td>
<td valign="middle" align="left">28.85&#xa0;&#xb1;&#xa0;0.93 a</td>
<td valign="middle" align="left">35.28&#xa0;&#xb1;&#xa0;1.56 ab</td>
</tr>
<tr>
<td valign="middle" align="center">OF</td>
<td valign="middle" align="left">0.20&#xa0;&#xb1;&#xa0;0.05 b</td>
<td valign="middle" align="left">2.11&#xa0;&#xb1;&#xa0;0.42 c</td>
<td valign="middle" align="left">34.62&#xa0;&#xb1;&#xa0;2.95 a</td>
<td valign="middle" align="left">29.50&#xa0;&#xb1;&#xa0;0.39 a</td>
<td valign="middle" align="left">33.57&#xa0;&#xb1;&#xa0;2.37 ab</td>
</tr>
<tr>
<td valign="middle" align="center">FA</td>
<td valign="middle" align="left">0.40&#xa0;&#xb1;&#xa0;0.06 a</td>
<td valign="middle" align="left">3.73&#xa0;&#xb1;&#xa0;0.54 b</td>
<td valign="middle" align="left">30.75&#xa0;&#xb1;&#xa0;1.80 a</td>
<td valign="middle" align="left">29.42&#xa0;&#xb1;&#xa0;1.41 a</td>
<td valign="middle" align="left">35.70&#xa0;&#xb1;&#xa0;0.61 a</td>
</tr>
<tr>
<td valign="middle" align="center">QL</td>
<td valign="middle" align="left">0.35&#xa0;&#xb1;&#xa0;0.08 a</td>
<td valign="middle" align="left">5.34&#xa0;&#xb1;&#xa0;0.72 a</td>
<td valign="middle" align="left">32.01&#xa0;&#xb1;&#xa0;1.85 a</td>
<td valign="middle" align="left">29.66&#xa0;&#xb1;&#xa0;3.25a</td>
<td valign="middle" align="left">32.64&#xa0;&#xb1;&#xa0;2.18 ab</td>
</tr>
<tr>
<td valign="middle" align="center">BC</td>
<td valign="middle" align="left">0.41&#xa0;&#xb1;&#xa0;0.04 a</td>
<td valign="middle" align="left">3.20&#xa0;&#xb1;&#xa0;0.08 b</td>
<td valign="middle" align="left">32.09&#xa0;&#xb1;&#xa0;2.93 a</td>
<td valign="middle" align="left">32.41&#xa0;&#xb1;&#xa0;2.58 a</td>
<td valign="middle" align="left">31.88&#xa0;&#xb1;&#xa0;0.93 b</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>Different lowercase letters within the same batch and selenium form indicate significant differences among treatments (<italic>P</italic>&#xa0;&lt;&#xa0;0.05).</p></fn>
</table-wrap-foot>
</table-wrap>
<p>After the second maize harvest, soil conditioners significantly affected soluble, exchangeable, and organically bound selenium, while iron-manganese-bound and residual selenium remained unaffected (<xref ref-type="table" rid="T2"><bold>Table&#xa0;2</bold></xref>). Biochar and lime significantly increased soluble selenium, whereas potassium humate and organic fertilizer treatments were comparable to CK. Exchangeable selenium increased significantly in the biochar treatment, but decreased under lime, potassium humate, and organic fertilizer compared to CK. Organically bound selenium did not differ significantly from CK, though organic fertilizer treatment exceeded biochar. Available selenium decreased in organic fertilizer, potassium humate, and lime treatments but increased in the biochar treatment relative to CK.</p>
<p>After the third maize batch, soil conditioners significantly influenced soluble, exchangeable, and residual selenium, but not iron-manganese-bound or organically bound selenium (<xref ref-type="table" rid="T2"><bold>Table&#xa0;2</bold></xref>). Biochar, lime, and potassium humate increased soluble selenium; lime also enhanced exchangeable selenium, while biochar and potassium humate increased it by 2.14 and 1.61 percentage points, respectively. Exchangeable and soluble selenium under organic fertilizer did not differ from CK. Residual selenium was significantly higher in potassium humate than in biochar, while CK values were comparable across treatments. Available selenium decreased by 0.57 percentage points under organic fertilizer, but increased by 1.25, 2.81, and 0.74 percentage points under potassium humate, lime, and biochar, respectively.</p>
<p>Overall, soil selenium continuously transforms among its forms. Iron-manganese-bound and residual selenium vary significantly between batches, reflecting dynamic soil-environment interactions. Despite these transformations, soluble and exchangeable selenium, which are directly available to crops, remain consistently low.</p>
</sec>
<sec id="s3_3">
<label>3.3</label>
<title>Maize growth characteristics under different soil conditioning measures</title>
<p>Total biomass of the first maize batch followed the order: biochar &gt; organic fertilizer &gt; CK &gt; potassium humate &gt; lime (<xref ref-type="fig" rid="f2"><bold>Figure&#xa0;2</bold></xref>). Compared to CK, biochar increased total biomass by 7.89%, whereas lime reduced it by 41.17%. Biochar notably enhanced root biomass by 30.20%, while lime reduced root biomass by 18.54%. Stem and leaf biomass did not differ significantly among biochar, potassium humate, organic fertilizer, and CK; however, lime significantly decreased stem and leaf biomass by 46.39%.</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Biomass of consecutively planted maize under different soil conditioner treatments. Different lowercase letters within the same batch indicate significant differences in root biomass among treatments (<italic>P</italic> &lt;&#xa0;0.05); different uppercase letters indicate significant differences in stem and leaf biomass among treatments (<italic>P</italic> &lt;&#xa0;0.05).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-17-1776194-g002.tif">
<alt-text content-type="machine-generated">Stacked bar chart comparing biomass dry weight per plant for roots (blue) and leaves and stems (green) across three batches and five treatments. Treatments differ significantly, as shown by letter groupings above and within each bar.</alt-text>
</graphic></fig>
<p>After the second maize batch, total biomass ranked as potassium humate &gt; organic fertilizer &gt; CK &gt; biochar &gt; lime (<xref ref-type="fig" rid="f2"><bold>Figure&#xa0;2</bold></xref>). Root biomass did not differ significantly among most treatments and the control, although potassium humate roots were significantly heavier than those under biochar and lime. Stem and leaf biomass was highest under potassium humate, increasing 25.85% compared to CK, whereas lime reduced stem and leaf biomass by 18.29%. Organic fertilizer and biochar had no significant effect on stem and leaf biomass relative to CK.</p>
<p>Following the third maize batch, total biomass followed the order: potassium humate &gt; organic fertilizer &gt; biochar &gt; CK &gt; lime (<xref ref-type="fig" rid="f2"><bold>Figure&#xa0;2</bold></xref>). Total biomass increased by 9.04%, 4.42%, and 2.54% under potassium humate, organic fertilizer, and biochar, respectively, relative to CK, whereas lime reduced total biomass by 19.42%. Potassium humate significantly enhanced root biomass by 29.53%, while other conditioners showed no significant difference compared to CK. Lime treatment caused a 21.42% reduction in stem and leaf biomass, whereas the other treatments were slightly higher than CK but not significantly different. Across the three maize batches, lime consistently inhibited growth. Organic fertilizers, potassium humate, and biochar exerted modest growth-promoting effects that became more apparent across consecutive plantings, with potassium humate showing the strongest positive impact on maize biomass.</p>
</sec>
<sec id="s3_4">
<label>3.4</label>
<title>Effects of soil conditioner applications on maize selenium nutrition</title>
<sec id="s3_4_1">
<label>3.4.1</label>
<title>Effects of soil conditioners on selenium content in maize</title>
<p>Soil conditioners altered selenium uptake and partitioning in maize, with distinct effects on roots versus aboveground tissues (<xref ref-type="fig" rid="f3"><bold>Figure&#xa0;3</bold></xref>). In roots, selenium content in nearly all conditioner treatments&#x2014;across all batches&#x2014;was higher than in the conventional treatment (CK), except for the first batch under potassium humate (<xref ref-type="fig" rid="f3"><bold>Figure&#xa0;3</bold></xref>). Lime and biochar significantly increased root selenium content relative to CK, and both were significantly higher than the organic fertilizer and potassium humate treatments; lime was also significantly higher than biochar. No significant differences were observed among CK, organic fertilizer, and potassium humate.</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Selenium content in maize roots and stems/leaves under different soil conditioning treatments. Different lowercase letters within the same batch column indicate significant differences among treatments (<italic>P</italic>&#xa0;&lt;&#xa0;0.05).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-17-1776194-g003.tif">
<alt-text content-type="machine-generated">Bar charts showing selenium contents in roots (left) and in leaves and stems (right) for three batches, comparing treatments CK, OF, FA, QL, and BC. QL shows the highest selenium content in both roots and aboveground parts, especially in batch one. Statistical differences among groups are indicated by letter annotations above each bar.</alt-text>
</graphic></fig>
<p>In stems and leaves, selenium content was generally lower under soil conditioner treatments than CK, except for the lime treatment and the third batch under biochar (<xref ref-type="fig" rid="f3"><bold>Figure&#xa0;3</bold></xref>). Selenium content in stems and leaves under lime increased by 87.38%, 40.13%, and 77.71% across the first to third batches, respectively, relative to CK. In contrast, organic fertilizer lowered selenium content by 13.67%, 48.85%, and 25.31% across the three batches, with significant decreases in the latter two. Potassium humate reduced stem and leaf selenium content by 8.48%, 23.15%, and 16.78% across the three batches, with a significant decrease in the second batch. Biochar reduced selenium content by 2.31% and 13.88% in the first two batches, but produced a slight (non-significant) increase of 5.60% in the third batch.</p>
<p>These findings indicate that while several soil conditioners enhance selenium absorption by roots, translocation of root-absorbed selenium to aboveground tissues remains limited. Differences in activation dynamics among conditioners are also evident, with biochar likely requiring longer periods to exert its effects.</p>
</sec>
<sec id="s3_4_2">
<label>3.4.2</label>
<title>Effects of conditioners on selenium accumulation in maize</title>
<p>Selenium accumulation patterns across consecutive maize batches are shown in <xref ref-type="fig" rid="f4"><bold>Figure&#xa0;4</bold></xref>. After the first harvest, conditioner treatments significantly affected selenium accumulation in roots but not in stems and leaves. Lime and biochar produced significantly higher root selenium accumulation than potassium humate, organic fertilizer, and CK, resulting in overall increases in total plant selenium accumulation of 38.17% and 35.03%, respectively, relative to CK.</p>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Selenium accumulation in maize under different soil conditioning treatments. Different lowercase letters within the same batch column indicate significant differences in root selenium accumulation (<italic>P</italic>&#xa0;&lt;&#xa0;0.05); different uppercase letters indicate significant differences in selenium accumulation in stems and leaves (<italic>P</italic>&#xa0;&lt;&#xa0;0.05).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-17-1776194-g004.tif">
<alt-text content-type="machine-generated">Stacked bar chart comparing selenium accumulation in roots and leaves and stems for three plant batches under different treatments (CK, OF, FA, QL, BC). Roots are orange, leaves and stems green. Batch 1 shows highest overall accumulations, particularly for QL and BC, with decreasing values in Batches 2 and 3. Error bars and statistical labels indicate significant differences among treatments and plant parts.</alt-text>
</graphic></fig>
<p>By the second batch, soil conditioners significantly influenced selenium accumulation in both roots and stems/leaves. Root selenium accumulation followed the order potassium humate &gt; biochar &gt; lime &gt; organic fertilizer &gt; CK. Compared to CK, selenium accumulation increased by 58.25%, 39.04%, 27.62%, and 0.49% in the potassium humate, biochar, lime, and organic fertilizer treatments, respectively, with significant increases under potassium humate and biochar. Stem and leaf selenium accumulation ranked as lime &gt; CK &gt; potassium humate &gt; biochar &gt; organic fertilizer, with the organic fertilizer treatment significantly lower than CK. Total plant selenium accumulation increased by 19.02% under lime and 15.97% under potassium humate, while biochar and organic fertilizer decreased accumulation by 1.28% and 32.36%, respectively.</p>
<p>In the third batch, biochar produced the highest root selenium accumulation, followed by potassium humate; both significantly exceeded CK. Lime and organic fertilizer also increased root selenium accumulation, although not significantly. In stems and leaves, lime again showed the strongest effect, significantly surpassing CK and all other treatments, whereas the remaining treatments showed no significant differences from CK. Total plant selenium accumulation followed the order lime &gt; biochar &gt; potassium humate &gt; CK &gt; organic fertilizer. Relative to CK, lime, biochar, and potassium humate increased selenium accumulation by 32.75%, 26.61%, and 9.84%, respectively, whereas organic fertilizer reduced it by 7.62%.</p>
</sec>
<sec id="s3_4_3">
<label>3.4.3</label>
<title>Effects of soil conditioners on the selenium transfer coefficient in maize</title>
<p>Nutrient uptake in plants typically proceeds from the roots to aboveground tissues, and the transfer coefficient reflects the efficiency with which absorbed nutrients are translocated internally. <xref ref-type="table" rid="T3"><bold>Table&#xa0;3</bold></xref> summarizes the selenium transfer coefficient from roots to shoots under different soil conditioner treatments. Both the type of conditioner and the time elapsed after application markedly influenced selenium translocation.</p>
<table-wrap id="T3" position="float">
<label>Table&#xa0;3</label>
<caption>
<p>Selenium transfer coefficient (TI) in maize plants under different treatments.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="center">Treatments</th>
<th valign="middle" align="center">Batch 1</th>
<th valign="middle" align="center">Batch 2</th>
<th valign="middle" align="center">Batch 3</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="center">CK</td>
<td valign="middle" align="center">0.3223&#xa0;&#xb1;&#xa0;0.0125 a</td>
<td valign="middle" align="center">0.4353&#xa0;&#xb1;&#xa0;0.0275 a</td>
<td valign="middle" align="center">0.1934&#xa0;&#xb1;&#xa0;0.0149 b</td>
</tr>
<tr>
<td valign="middle" align="center">OF</td>
<td valign="middle" align="center">0.2496&#xa0;&#xb1;&#xa0;0.0042 b</td>
<td valign="middle" align="center">0.2033&#xa0;&#xb1;&#xa0;0.0360 c</td>
<td valign="middle" align="center">0.1328&#xa0;&#xb1;&#xa0;0.0100 c</td>
</tr>
<tr>
<td valign="middle" align="center">FA</td>
<td valign="middle" align="center">0.3074&#xa0;&#xb1;&#xa0;0.0363 a</td>
<td valign="middle" align="center">0.2643&#xa0;&#xb1;&#xa0;0.0268 bc</td>
<td valign="middle" align="center">0.1482&#xa0;&#xb1;&#xa0;0.0146 c</td>
</tr>
<tr>
<td valign="middle" align="center">QL</td>
<td valign="middle" align="center">0.2573&#xa0;&#xb1;&#xa0;0.0311 b</td>
<td valign="middle" align="center">0.4283&#xa0;&#xb1;&#xa0;0.0762 a</td>
<td valign="middle" align="center">0.2587&#xa0;&#xb1;&#xa0;0.0160 a</td>
</tr>
<tr>
<td valign="middle" align="center">BC</td>
<td valign="middle" align="center">0.2230&#xa0;&#xb1;&#xa0;0.0120 b</td>
<td valign="middle" align="center">0.2906&#xa0;&#xb1;&#xa0;0.0052 b</td>
<td valign="middle" align="center">0.1539&#xa0;&#xb1;&#xa0;0.0106 c</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>TI, Transport Index = selenium content in stems and leaves/selenium content in roots. Different uppercase letters within the same batch indicate significant differences among treatments (<italic>P</italic>&#xa0;&lt;&#xa0;0.05).</p></fn>
</table-wrap-foot>
</table-wrap>
<p>In the first maize batch, all soil conditioners reduced the selenium transfer coefficient, with organic fertilizer, lime, and biochar causing the largest declines. By the second batch, the transfer coefficients under all conditioner treatments remained lower than CK, yet lime showed a pronounced increase, becoming statistically indistinguishable from CK and significantly higher than the other conditioners. In the third batch, lime exhibited a significantly higher selenium transfer coefficient than CK, whereas CK exceeded biochar, potassium humate, and organic fertilizer.</p>
<p>Across application cycles, the transfer coefficients under organic fertilizer and potassium humate consistently remained below CK and declined further over time, with organic fertilizer showing the steepest decrease. Biochar also consistently reduced the transfer coefficient relative to CK but displayed a fluctuating pattern&#x2014;first decreasing, then increasing, and finally decreasing again&#x2014;yielding values closer to CK than those observed under organic fertilizer or potassium humate. Lime reduced the transfer coefficient only in the first batch but exceeded CK thereafter, indicating that lime ultimately enhances selenium movement from roots to aboveground tissues over longer timescales.</p>
</sec>
</sec>
<sec id="s3_5">
<label>3.5</label>
<title>Interrelationships among factors influencing selenium absorption and translocation under different soil conditioners</title>
<p>Correlation analysis between maize selenium nutritional indicators and key soil physicochemical properties and selenium speciation is presented in <xref ref-type="table" rid="T4"><bold>Table&#xa0;4</bold></xref>. Among selenium fractions, soluble selenium showed significant or highly significant positive correlations with selenium content in stems and leaves, selenium accumulation in stems and leaves, and the selenium transfer coefficient. Iron&#x2013;manganese&#x2013;bound selenium exhibited highly significant positive correlations with root selenium content, stem and leaf selenium content, selenium accumulation in both roots and shoots, total plant selenium accumulation, and the selenium transfer coefficient. In contrast, residual selenium was highly negatively correlated with all of these indicators. Exchangeable and organically bound selenium showed no significant correlations with maize selenium nutritional parameters.</p>
<table-wrap id="T4" position="float">
<label>Table&#xa0;4</label>
<caption>
<p>Linear correlation coefficients between soil properties, soil selenium fractions, and maize selenium nutritional indicators.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left"/>
<th valign="middle" align="center">Se content in roots</th>
<th valign="middle" align="center">Se content in leaves and stems</th>
<th valign="middle" align="center">Se accumulation in roots</th>
<th valign="middle" align="center">Se accumulation in straws</th>
<th valign="middle" align="center">Total Se accumulation in plants</th>
<th valign="middle" align="center">Transport indexes</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="left">Soluble- Se</td>
<td valign="middle" align="center">0.172</td>
<td valign="middle" align="center"><bold>0.350<sup>**</sup></bold></td>
<td valign="middle" align="center">0.153</td>
<td valign="middle" align="center"><bold>0.299<sup>*</sup></bold></td>
<td valign="middle" align="center">0.228</td>
<td valign="middle" align="center"><bold>0.431<sup>**</sup></bold></td>
</tr>
<tr>
<td valign="middle" align="left">Exchangeable- Se</td>
<td valign="middle" align="center">-0.098</td>
<td valign="middle" align="center">-0.186</td>
<td valign="middle" align="center">-0.130</td>
<td valign="middle" align="center">-0.242</td>
<td valign="middle" align="center">-0.188</td>
<td valign="middle" align="center">-0.181</td>
</tr>
<tr>
<td valign="middle" align="left">Fe/Mn oxide-bound-Se</td>
<td valign="middle" align="center"><bold>0.565<sup>**</sup></bold></td>
<td valign="middle" align="center"><bold>0.707<sup>**</sup></bold></td>
<td valign="middle" align="center"><bold>0.616<sup>**</sup></bold></td>
<td valign="middle" align="center"><bold>0.797<sup>**</sup></bold></td>
<td valign="middle" align="center"><bold>0.732<sup>**</sup></bold></td>
<td valign="middle" align="center"><bold>0.468<sup>**</sup></bold></td>
</tr>
<tr>
<td valign="middle" align="left">Organic Matter-bound-Se</td>
<td valign="middle" align="center">0.060</td>
<td valign="middle" align="center">0.139</td>
<td valign="middle" align="center">0.051</td>
<td valign="middle" align="center">0.147</td>
<td valign="middle" align="center">0.097</td>
<td valign="middle" align="center">0.173</td>
</tr>
<tr>
<td valign="middle" align="left">Residual-Se</td>
<td valign="middle" align="center"><bold>-0.470<sup>**</sup></bold></td>
<td valign="middle" align="center"><bold>-0.667<sup>**</sup></bold></td>
<td valign="middle" align="center"><bold>-0.462<sup>**</sup></bold></td>
<td valign="middle" align="center"><bold>-0.687<sup>**</sup></bold></td>
<td valign="middle" align="center"><bold>-0.590<sup>**</sup></bold></td>
<td valign="middle" align="center"><bold>-0.559<sup>**</sup></bold></td>
</tr>
<tr>
<td valign="middle" align="left">pH</td>
<td valign="middle" align="center"><bold>0.504<sup>**</sup></bold></td>
<td valign="middle" align="center"><bold>0.633<sup>**</sup></bold></td>
<td valign="middle" align="center"><bold>0.324<sup>*</sup></bold></td>
<td valign="middle" align="center"><bold>0.380<sup>**</sup></bold></td>
<td valign="middle" align="center"><bold>0.367<sup>**</sup></bold></td>
<td valign="middle" align="center"><bold>0.411<sup>**</sup></bold></td>
</tr>
<tr>
<td valign="middle" align="left">Available P</td>
<td valign="middle" align="center">0.203</td>
<td valign="middle" align="center">0.173</td>
<td valign="middle" align="center">-0.007</td>
<td valign="middle" align="center">-0.131</td>
<td valign="middle" align="center">-0.063</td>
<td valign="middle" align="center">-0.056</td>
</tr>
<tr>
<td valign="middle" align="left">Available S</td>
<td valign="middle" align="center">-0.032</td>
<td valign="middle" align="center">-0.162</td>
<td valign="middle" align="center">0.029</td>
<td valign="middle" align="center">-0.073</td>
<td valign="middle" align="center">-0.016</td>
<td valign="middle" align="center"><bold>-0.290<sup>*</sup></bold></td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>*indicates <italic>P</italic> &lt;&#xa0;0.05, **indicates <italic>P</italic>&#xa0;&lt;&#xa0;0.01.</p></fn>
</table-wrap-foot>
</table-wrap>
<p>Regarding soil properties, soil pH displayed significant or highly significant positive correlations with root selenium content, stem and leaf selenium content, selenium accumulation in roots and shoots, total selenium accumulation, and the selenium transfer coefficient. Among available nutrients, available sulfur showed a significant negative correlation only with the selenium transfer coefficient, whereas available phosphorus displayed no significant associations.</p>
<p>Overall, these findings indicate that soil pH is a key environmental factor regulating maize selenium nutrition under soil conditioner treatments. Soluble selenium, iron&#x2013;manganese&#x2013;bound selenium, and residual selenium serve as critical indicators of selenium uptake and accumulation, while available sulfur significantly influences selenium translocation within the plant.</p>
<p><xref ref-type="fig" rid="f5"><bold>Figure&#xa0;5</bold></xref> shows principal component analysis (PCA) of the three batches of maize data with added soil conditioner. The results show that the first two principal components (PC1 and PC2) cumulatively explained 60.5%, 60.1%, and 59.9% of the total variance, respectively, indicating that these two dimensions can reflect the original structure of the data well. In the first batch of maize, the lime application treatment showed a significant difference from the control (CK) treatment. Soil pH, available phosphorus in the soil, and selenium content in different parts of the maize plant were closely related to lime application. In the second and third batches of maize planting, there was significant overlap among the treatments, indicating a weakening of the treatment effect. In the second batch of maize, soluble selenium content, transfer coefficient, selenium content in maize stems and leaves, and soil pH were closely related; selenium content in maize roots was closely related to available phosphorus in the soil. In the third batch of maize, the selenium content in both maize roots and stems and leaves was closely related to exchangeable selenium, while the selenium content in maize stems and leaves, soil pH, and selenium transfer coefficient were closely related.</p>
<fig id="f5" position="float">
<label>Figure&#xa0;5</label>
<caption>
<p>Principal Component Analysis of soil selenium speciation, maize biomass, and selenium yield under soil conditioner addition. CK, OF, FA, QL, and BC represent control, organic fertilizer, potassium humate, lime, and biochar pretreatment, respectively. Sol-Se, Exc-Se, FMO-Se, OM-Se, and Res-Se represent Soluble Se, Exchangeable Fe/Mn oxide-bound Se, Organic matter-bond Se, and Residual Se, respectively. AP and AS represent available phosphorus and available sulfur, respectively. Biomass-roots and Biomass-SL represent the biomass of maize roots and stems/leafs, respectively. Se-roots and Se-SL represent the selenium content of maize roots and stems/leafs, respectively. TI represents the selenium transfer coefficient.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-17-1776194-g005.tif">
<alt-text content-type="machine-generated">Principal component analysis biplots for three batches display relationships among treatments (CK, OF, FA, QL, BC) and environmental variables, with distinct colored ellipses and vectors for each variable, highlighting differences in variance explained by PC1 and PC2.</alt-text>
</graphic></fig>
</sec>
</sec>
<sec id="s4" sec-type="discussion">
<label>4</label>
<title>Discussion</title>
<sec id="s4_1">
<label>4.1</label>
<title>Effects of soil conditioners on soil selenium species</title>
<p>This study continuously monitored shifts in soil selenium speciation across three sequential maize seedling harvests and found substantial variation in several selenium fractions among batches under the same treatment, particularly in iron&#x2013;manganese-bound and residual selenium (<xref ref-type="table" rid="T1"><bold>Table&#xa0;1</bold></xref>). Soil pH, available phosphorus, and available sulfur are key factors influencing soil selenium speciation and crop selenium uptake. These fluctuations were likely driven not solely by the applied soil conditioners but also by interacting climatic, hydrologic, and crop-management factors. Research on arsenic speciation indicates that arsenic is initially adsorbed onto iron&#x2013;aluminum clay mineral surfaces and, as soil pH fluctuates, these minerals transform and incorporate the adsorbed arsenic into their lattices, resulting in more stable fixation (<xref ref-type="bibr" rid="B28">Ran, 2023</xref>). Given the structural and chemical similarities between selenium and arsenic, iron&#x2013;manganese-bound and residual selenium likely undergo comparable inter-conversion during clay-mineral transformations under variable environmental conditions.</p>
<p>Soil conditioners significantly influenced exchangeable selenium after all three maize harvests; soluble selenium after the second and third harvests; residual selenium during the first and third harvests; and organically bound selenium after the second harvest (<xref ref-type="table" rid="T1"><bold>Table&#xa0;1</bold></xref>). Soluble and exchangeable selenium represent the bioavailable fractions in soil, and their availability is strongly regulated by soil pH and organic matter content (<xref ref-type="bibr" rid="B19">Liu et&#xa0;al., 2015</xref>). The addition of soil conditioners markedly altered soil pH (<xref ref-type="fig" rid="f1"><bold>Figure&#xa0;1</bold></xref>), with treatments containing organic fertilizer or potassium humate producing the lowest pH values and correspondingly the lowest available selenium.</p>
<p>The four soil conditioners used in this study differed substantially in organic-matter characteristics, which influenced selenium sorption and stabilization. Because selenium fixation by organic matter requires time to develop, significant differences in organically bound selenium among treatments did not emerge until the second harvest. Organic fertilizer and potassium humate, both rich in active organic components, produced higher levels of organically bound selenium at that stage. Biochar, characterized by a more stable carbon matrix, exerted a delayed effect, displaying elevated organically bound selenium only after the third maize harvest.</p>
<p>The observed effects of soil conditioners on residual selenium were consistent with previous findings (<xref ref-type="bibr" rid="B32">Tan et&#xa0;al., 2024</xref>; <xref ref-type="bibr" rid="B21">Lu et&#xa0;al., 2023</xref>). Residual selenium, immobilized within mineral lattices, is largely inaccessible under stable natural conditions. During the first planting cycle, the combined initial activation from irrigation, maize cultivation, and soil-conditioner application likely disrupted lattice structures, leading to treatment-specific alterations in residual selenium. The pronounced differences among conditioners consequently produced varying degrees of influence on this most recalcitrant selenium fraction.</p>
</sec>
<sec id="s4_2">
<label>4.2</label>
<title>Effects of soil conditioners on selenium absorption and accumulation in maize</title>
<p>The application of soil conditioners increased selenium concentrations in maize roots, though the magnitude of this increase varied markedly among treatments (<xref ref-type="fig" rid="f3"><bold>Figure&#xa0;3</bold></xref>). This pattern is consistent with previous findings. For example, <xref ref-type="bibr" rid="B15">Huang et&#xa0;al. (2019)</xref> reported that selenium uptake by rice roots differed substantially depending on how sodium selenite was combined with various organic materials, while <xref ref-type="bibr" rid="B23">Ma et&#xa0;al. (2023)</xref> found that biochar application in dry-land systems enhanced plant selenium absorption. These differences arise because soil conditioners modify soil physicochemical properties and selenium speciation, influencing plant uptake of distinct selenium chemical forms (<xref ref-type="bibr" rid="B6">Dinh et&#xa0;al., 2019</xref>).</p>
<p>Lime application significantly elevated root selenium content, largely due to its strong alkalizing effect on soil pH (<xref ref-type="fig" rid="f1"><bold>Figure&#xa0;1</bold></xref>). Soil pH showed a highly significant positive correlation with selenium content in both roots and leaves (<xref ref-type="table" rid="T4"><bold>Table&#xa0;4</bold></xref>), consistent with earlier studies (<xref ref-type="bibr" rid="B21">Lu et&#xa0;al., 2023</xref>). Although all conditioners increased root selenium, the selenium content in the aboveground tissues of maize treated with organic fertilizer, potassium humate, or biochar was lower than that of the control (<xref ref-type="fig" rid="f3"><bold>Figure&#xa0;3</bold></xref>). This suggests that selenium absorbed in these treatments was not effectively translocated from the roots to shoots. Excessive selenium (Se) acts as a pro-oxidant in plants, causing significant toxicity (selenosis) characterized by stunted growth, root inhibition, chlorosis, and necrosis (<xref ref-type="bibr" rid="B11">Gu et&#xa0;al., 2025</xref>). It causes damage by replacing sulfur in amino acids (cysteine and methionine), resulting in dysfunctional proteins, alongside inducing severe oxidative stress (<xref ref-type="bibr" rid="B2">Cao et&#xa0;al., 2025</xref>).</p>
<p>Crops can absorb all available selenium species in soil, including Se(IV), Se(VI), and dissolved organic selenium. However, selenium must undergo intracellular transformation before being transported to aboveground tissues. Se(IV) taken up by roots is quickly converted into organoselenium compounds primarily selenomethionine and methylselenocysteine most of which remain in the roots, with only limited translocation to stems (<xref ref-type="bibr" rid="B37">Winkel et&#xa0;al., 2015</xref>). In contrast, selenate (Se(VI)) is readily transported through the xylem to aboveground tissues, where it is reduced to Se(IV); following reduction, selenium is incorporated into organoselenium forms and redistributed to various tissues through both xylem and phloem pathways [32]. Transport efficiencies differ widely across chemical forms, with the general order Se(VI) &gt; SeMet &gt; Se(IV)/SeCys (<xref ref-type="bibr" rid="B16">Kikkert and Berkelaar, 2013</xref>).</p>
<p>Different soil selenium fractions also contain distinct chemical species (<xref ref-type="bibr" rid="B27">Qu et&#xa0;al., 1997</xref>). Soluble selenium is dominated by SeO<sub>4</sub>&#xb2;<sup>-</sup>, with smaller proportions of SeO<sub>3</sub>&#xb2;<sup>-</sup>and soluble organic selenium, whereas exchangeable selenium exists mainly as SeO<sub>3</sub>&#xb2;<sup>-</sup>(<xref ref-type="bibr" rid="B10">Gissel et&#xa0;al., 1984</xref>; <xref ref-type="bibr" rid="B35">White, 2016</xref>). In this study, soil conditioners significantly affected exchangeable selenium (<xref ref-type="table" rid="T1"><bold>Table&#xa0;1</bold></xref>), and selenium content and transfer coefficients in stems and leaves were significantly positively correlated with soluble selenium (<xref ref-type="table" rid="T4"><bold>Table&#xa0;4</bold></xref>). This indicates that the application of soil conditioners on dryland may mainly affect the content of Se(IV) in available selenium in the soil, thereby affecting the absorption and accumulation of selenium by crops.</p>
<p>Furthermore, plant absorption of selenium involves complex synergistic and competitive relationships with other nutrients. Generally, Se(IV) in the soil is absorbed by phosphorus transport proteins (<xref ref-type="bibr" rid="B42">Zhang et&#xa0;al., 2014</xref>), while Se(VI) mainly enters the plant root system through sulfur transport proteins (<xref ref-type="bibr" rid="B12">Gupta and Gupta, 2017</xref>). Therefore, the availability of phosphorus and sulfur in the soil plays an important regulatory role in the efficiency of selenium translocation, absorption, and accumulation in plants. Experiments have shown that the combined application of phosphorus and Se(IV) significantly reduces selenium translocation and accumulation in wheat, but the combined application of phosphorus and Se(VI) increases selenium absorption and accumulation in wheat (<xref ref-type="bibr" rid="B41">Zhang et&#xa0;al., 2017</xref>). The effect of sulfur on plant selenium absorption and accumulation also varies depending on the crop&#x2019;s sulfur status (<xref ref-type="bibr" rid="B31">Stroud et&#xa0;al., 2010</xref>). The addition of soil conditioners also significantly affects the transformation characteristics of available phosphorus, available sulfur, and selenium forms in the soil, thereby affecting the absorption and translocation of selenium by maize plants. However, as mentioned above, we regret that we did not detect changes in soil Se(IV) and Se(VI). Therefore, we believe that future related studies must measure changes in soil Se(IV) and Se(VI) content to help understand the mechanism of selenium transfer and transformation in soil and plants.</p>
</sec>
</sec>
<sec id="s5" sec-type="conclusions">
<label>5</label>
<title>Conclusions</title>
<p>Different soil conditioners had significant effects on soil selenium speciation. Biochar consistently increased both soluble and exchangeable selenium, whereas lime enhanced soluble selenium but significantly reduced exchangeable selenium after the second harvest compared with the control. All conditioners increased root selenium concentrations, with lime and biochar producing the most pronounced effects. However, except for lime, treatments reduced aboveground selenium content relative to the control, with organic fertilizer and potassium humate causing the greatest declines. Transfer coefficients in all conditioner treatments were lower than those of the control, reflecting their substantial influence on residual selenium. Correlation analysis showed that root selenium, aboveground selenium, and transfer coefficients were all significantly positively correlated with soil pH but not with exchangeable selenium. Soluble selenium content and transfer coefficients were also significantly positively correlated with soil soluble selenium. Overall, applying soil conditioners to selenium-rich dry-land soils increased soil pH and enhanced available selenium, thereby improving selenium uptake by maize roots. However, the selenium forms enhanced by conditioners were less efficiently transported to aboveground tissues. Among the treatments, lime was the most effective at increasing crop selenium content, while biochar was most effective at enhancing soil available selenium. The following aspects should be emphasized the limitations of the study and future studies: Techniques are necessary to develop and utilize such as DGT to improve the accuracy of the evaluation of Se bioavailability. Studies are required to acquire understanding about antagonistic and competitive effects of each nutrient elements (N, P, and S) with Se on plant uptake of Se. Studies are needed to explore the role, as well as the strength of each component (e.g., LMWOAs, HA, FA) in organic materials on Se bioavailability. Models are needed to predict the effects of climate change on the soil distribution of available Se on a large-scale.</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 authors.</p></sec>
<sec id="s7" sec-type="author-contributions">
<title>Author contributions</title>
<p>YH: Conceptualization, Funding acquisition, Project administration, Supervision, Writing &#x2013; original draft. TH: Data curation, Software, Visualization, Writing &#x2013; review &amp; editing. SL: Data curation, Software, Validation, Visualization, Writing &#x2013; review &amp; editing. ML: Conceptualization, Data curation, Writing &#x2013; review &amp; editing. JW: Project administration, Resources, Software, Writing &#x2013; review &amp; editing. XP: Investigation, Methodology, Writing &#x2013; review &amp; editing. CW: Formal analysis, Software, Writing &#x2013; review &amp; editing. ZL: Investigation, Software, Writing &#x2013; review &amp; editing. BL: Project administration, Resources, Writing &#x2013; review &amp; editing. QT: Formal analysis, Investigation, Validation, Writing &#x2013; review &amp; editing. ZJ: Conceptualization, Formal analysis, Funding acquisition, Writing &#x2013; review &amp; editing.</p></sec>
<sec id="s9" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The author(s) declared that this work 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) declared that generative AI was not used in the creation of this manuscript.</p>
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<ref-list>
<title>References</title>
<ref id="B1">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>An</surname> <given-names>J.</given-names></name>
<name><surname>Zhang</surname> <given-names>D.</given-names></name>
<name><surname>Ye</surname> <given-names>J.</given-names></name>
<name><surname>Luo</surname> <given-names>D.</given-names></name>
<name><surname>Wang</surname> <given-names>P.</given-names></name>
</person-group> (<year>2019</year>). 
<article-title>Effects of different phosphorus sources on soil selenium speciation and selenium absorption and translocation in wheat</article-title>. <source>Jiangsu Agric. Sci.</source> <volume>47</volume>, <fpage>119</fpage>&#x2013;<lpage>122</lpage>.
</mixed-citation>
</ref>
<ref id="B2">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Cao</surname> <given-names>H.</given-names></name>
<name><surname>Xiao</surname> <given-names>X.</given-names></name>
<name><surname>Zhang</surname> <given-names>X.</given-names></name>
<name><surname>Yu</surname> <given-names>L.</given-names></name>
</person-group> (<year>2025</year>). 
<article-title>The bio-applications of low-valence-selenium-substituted glucoses</article-title>. <source>Chin. Chem. Lett.</source>, <fpage>112197</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.cclet.2025.112197</pub-id>, PMID: <pub-id pub-id-type="pmid">41760527</pub-id>
</mixed-citation>
</ref>
<ref id="B3">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Daud</surname> <given-names>M. F. B.</given-names></name>
<name><surname>Rempelos</surname> <given-names>L.</given-names></name>
<name><surname>Cakmak</surname> <given-names>I.</given-names></name>
<name><surname>Leifert</surname> <given-names>C.</given-names></name>
<name><surname>Bilsborrow</surname> <given-names>P.</given-names></name>
</person-group> (<year>2024</year>). 
<article-title>Increasing grain selenium concentration via genetic and agronomic innovations</article-title>. <source>Plant Soil</source> <volume>494</volume>, <fpage>477</fpage>&#x2013;<lpage>496</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/s11104-023-06293-z</pub-id>, PMID: <pub-id pub-id-type="pmid">41758449</pub-id>
</mixed-citation>
</ref>
<ref id="B4">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Deng</surname> <given-names>X.</given-names></name>
<name><surname>Zhao</surname> <given-names>Z.</given-names></name>
<name><surname>Lv</surname> <given-names>C.</given-names></name>
<name><surname>Zhang</surname> <given-names>Z.</given-names></name>
<name><surname>Yuan</surname> <given-names>L.</given-names></name>
<name><surname>Liu</surname> <given-names>X.</given-names></name>
</person-group> (<year>2021</year>). 
<article-title>Effects of sulfur application on selenium uptake and seed selenium speciation in soybean (<italic>Glycine max</italic> L.) grown in different soil types</article-title>. <source>Ecotoxicology Environ. Saf.</source> <volume>209</volume>, <fpage>111790</fpage>.
</mixed-citation>
</ref>
<ref id="B5">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Dinh</surname> <given-names>Q. T.</given-names></name>
<name><surname>Cui</surname> <given-names>Z.</given-names></name>
<name><surname>Huang</surname> <given-names>J.</given-names></name>
<name><surname>Tran</surname> <given-names>T. A. T.</given-names></name>
<name><surname>Wang</surname> <given-names>D.</given-names></name>
<name><surname>Yang</surname> <given-names>W.</given-names></name>
<etal/>
</person-group>. (<year>2018</year>). 
<article-title>Selenium distribution in the Chinese environment and its relationship with human health: a review</article-title>. <source>Environ. Int.</source> <volume>112</volume>, <fpage>294</fpage>&#x2013;<lpage>309</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.envint.2017.12.035</pub-id>, PMID: <pub-id pub-id-type="pmid">29438838</pub-id>
</mixed-citation>
</ref>
<ref id="B6">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Dinh</surname> <given-names>Q. T.</given-names></name>
<name><surname>Wang</surname> <given-names>M.</given-names></name>
<name><surname>Tran</surname> <given-names>T. A. T.</given-names></name>
<name><surname>Zhou</surname> <given-names>F.</given-names></name>
<name><surname>Wang</surname> <given-names>D.</given-names></name>
<name><surname>Zhai</surname> <given-names>H.</given-names></name>
<etal/>
</person-group>. (<year>2019</year>). 
<article-title>Bioavailability of selenium in soil-plant system and a regulatory approach</article-title>. <source>Crit. Rev. Environ. Sci. Technol.</source> <volume>49</volume>, <fpage>443</fpage>&#x2013;<lpage>517</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1080/10643389.2018.1550987</pub-id>, PMID: <pub-id pub-id-type="pmid">41735180</pub-id>
</mixed-citation>
</ref>
<ref id="B7">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Du</surname> <given-names>Y.</given-names></name>
<name><surname>Luo</surname> <given-names>K.</given-names></name>
<name><surname>Ni</surname> <given-names>R.</given-names></name>
<name><surname>Hussain</surname> <given-names>R.</given-names></name>
</person-group> (<year>2018</year>). 
<article-title>Selenium and hazardous elements distribution in plant-soil-water system and human health risk assessment of lower Cambrian, Southern Shaanxi, China</article-title>. <source>Environ. Geochemistry Health</source> <volume>40</volume>, <fpage>2049</fpage>&#x2013;<lpage>2069</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/s10653-018-0082-3</pub-id>, PMID: <pub-id pub-id-type="pmid">29497886</pub-id>
</mixed-citation>
</ref>
<ref id="B8">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Farooq</surname> <given-names>M. R.</given-names></name>
<name><surname>Zhang</surname> <given-names>Z. Z.</given-names></name>
<name><surname>Yuan</surname> <given-names>L. X.</given-names></name>
<name><surname>Liu</surname> <given-names>X. D.</given-names></name>
<name><surname>Rehman</surname> <given-names>A.</given-names></name>
<name><surname>Ba&#xf1;uelos</surname> <given-names>G. S.</given-names></name>
<etal/>
</person-group>. (<year>2023</year>). 
<article-title>Influencing factors on bioavailability and spatial distribution of soil selenium in dry semi-arid area</article-title>. <source>Agriculture</source> <volume>13</volume>, <fpage>1</fpage>&#x2013;<lpage>17</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.3390/agriculture13030576</pub-id>, PMID: <pub-id pub-id-type="pmid">41725453</pub-id>
</mixed-citation>
</ref>
<ref id="B9">
<mixed-citation publication-type="book">
<person-group person-group-type="author">
<name><surname>Fordyce</surname> <given-names>F. M.</given-names></name>
</person-group> (<year>2013</year>). &#x201c;
<article-title>Selenium deficiency and toxicity in the environment</article-title>,&#x201d; in <source>Essentials of medical geology</source>. Eds. 
<person-group person-group-type="editor">
<name><surname>Selinus</surname> <given-names>O.</given-names></name>
<name><surname>Alloway</surname> <given-names>B.</given-names></name>
<name><surname>Centeno</surname> <given-names>J. A.</given-names></name>
<etal/>
</person-group> (
<publisher-name>Springer</publisher-name>, <publisher-loc>Dordrecht</publisher-loc>), <fpage>375</fpage>&#x2013;<lpage>416</lpage>.
</mixed-citation>
</ref>
<ref id="B10">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Gissel-Nielsen</surname> <given-names>G.</given-names></name>
<name><surname>Gupta</surname> <given-names>U. C.</given-names></name>
<name><surname>Lamand</surname> <given-names>M.</given-names></name>
<name><surname>Westermarck</surname> <given-names>T.</given-names></name>
</person-group> (<year>1984</year>). 
<article-title>Selenium in soil and plant and its importance in livestock and human nutrition</article-title>. <source>Adv. Agron.</source> <volume>37</volume>, <fpage>397</fpage>&#x2013;<lpage>460</lpage>.
</mixed-citation>
</ref>
<ref id="B11">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Gu</surname> <given-names>J.</given-names></name>
<name><surname>Gu</surname> <given-names>J.</given-names></name>
<name><surname>Yu</surname> <given-names>L.</given-names></name>
</person-group> (<year>2025</year>). 
<article-title>Selenium and alzheimer's disease</article-title>. <source>Chin. Chem. Letters.</source> <volume>36</volume>, <elocation-id>110727</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.cclet.2024.110727</pub-id>, PMID: <pub-id pub-id-type="pmid">41760527</pub-id>
</mixed-citation>
</ref>
<ref id="B12">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Gupta</surname> <given-names>M.</given-names></name>
<name><surname>Gupta</surname> <given-names>S.</given-names></name>
</person-group> (<year>2017</year>). 
<article-title>An overview of selenium uptake, metabolism, and toxicity in plants</article-title>. <source>Front. Plant Sci.</source> <volume>7</volume>, <elocation-id>2074</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.3389/fpls.2016.02074</pub-id>, PMID: <pub-id pub-id-type="pmid">28123395</pub-id>
</mixed-citation>
</ref>
<ref id="B13">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Haug</surname> <given-names>A.</given-names></name>
<name><surname>Graham</surname> <given-names>R. D.</given-names></name>
<name><surname>Christophersen</surname> <given-names>O. A.</given-names></name>
<name><surname>Lyons</surname> <given-names>G. H.</given-names></name>
</person-group> (<year>2007</year>). 
<article-title>How to use the world&#x2019;s scarce selenium resources efficiently to increase the selenium concentration in food</article-title>. <source>Microbial Ecol. Health Dis.</source> <volume>19</volume>, <fpage>209</fpage>&#x2013;<lpage>228</lpage>.
</mixed-citation>
</ref>
<ref id="B14">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Huang</surname> <given-names>J.</given-names></name>
<name><surname>Huang</surname> <given-names>X.</given-names></name>
<name><surname>Jiang</surname> <given-names>D.</given-names></name>
</person-group> (<year>2024</year>). 
<article-title>Phosphorus can effectively reduce selenium adsorption in selenium-rich lateritic red soil</article-title>. <source>Sci. Total Environ.</source> <volume>906</volume>, <fpage>167356</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.scitotenv.2023.167356</pub-id>, PMID: <pub-id pub-id-type="pmid">37769720</pub-id>
</mixed-citation>
</ref>
<ref id="B15">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Huang</surname> <given-names>T.</given-names></name>
<name><surname>Jiang</surname> <given-names>Z.</given-names></name>
<name><surname>Liao</surname> <given-names>Q.</given-names></name>
<name><surname>Xing</surname> <given-names>Y.</given-names></name>
<name><surname>Liang</surname> <given-names>P.</given-names></name>
</person-group> (<year>2019</year>). 
<article-title>Effects of exogenous selenium combined with different materials on selenium nutrition and selenium utilization in rice</article-title>. <source>Soil</source> <volume>51</volume>, <fpage>269</fpage>&#x2013;<lpage>278</lpage>.
</mixed-citation>
</ref>
<ref id="B16">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Kikkert</surname> <given-names>J.</given-names></name>
<name><surname>Berkelaar</surname> <given-names>E.</given-names></name>
</person-group> (<year>2013</year>). 
<article-title>Plant uptake and translocation of inorganic and organic forms of selenium</article-title>. <source>Arch. Environ. ContamToxicol.</source> <volume>65</volume>, <fpage>458</fpage>&#x2013;<lpage>465</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/s00244-013-9926-0</pub-id>, PMID: <pub-id pub-id-type="pmid">23793939</pub-id>
</mixed-citation>
</ref>
<ref id="B17">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Li</surname> <given-names>Z.</given-names></name>
<name><surname>Liang</surname> <given-names>D.</given-names></name>
<name><surname>Peng</surname> <given-names>Q.</given-names></name>
<name><surname>Cui</surname> <given-names>Z.</given-names></name>
<name><surname>Huang</surname> <given-names>J.</given-names></name>
<name><surname>Lin</surname> <given-names>Z.</given-names></name>
</person-group> (<year>2017</year>). 
<article-title>Interaction between selenium and soil organic matter and its impact on soil selenium bioavailability A review</article-title>. <source>Geoderma</source> <volume>295</volume>, <fpage>69</fpage>&#x2013;<lpage>79</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.geoderma.2017.02.019</pub-id>, PMID: <pub-id pub-id-type="pmid">41760527</pub-id>
</mixed-citation>
</ref>
<ref id="B18">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Liu</surname> <given-names>NN.</given-names></name>
<name><surname>Wang</surname> <given-names>M.</given-names></name>
<name><surname>Zhou</surname> <given-names>F.</given-names></name>
<name><surname>Zhai</surname> <given-names>H.</given-names></name>
<name><surname>Qi</surname> <given-names>MX.</given-names></name>
<name><surname>Liu</surname> <given-names>Y.</given-names></name>
<etal/>
</person-group>. (<year>2021</year>). 
<article-title>Selenium bioavailability in soil-wheat system and its dominant influential factors: A field study in Shaanxi Province, China</article-title>. <source>Sci. Total Environ.</source>, <fpage>770</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.scitotenv.2020.144664</pub-id>, PMID: <pub-id pub-id-type="pmid">33513517</pub-id>
</mixed-citation>
</ref>
<ref id="B19">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Liu</surname> <given-names>X.</given-names></name>
<name><surname>Zhao</surname> <given-names>Z.</given-names></name>
<name><surname>Duan</surname> <given-names>B.</given-names></name>
<name><surname>Hu</surname> <given-names>C.</given-names></name>
<name><surname>Zhao</surname> <given-names>X.</given-names></name>
<name><surname>Guo</surname> <given-names>Z.</given-names></name>
<etal/>
</person-group>. (<year>2015</year>). 
<article-title>Effect of applied sulphur on the uptake by wheat of selenium applied as selenite</article-title>. <source>Plant Soil</source> <volume>386</volume>, <fpage>35</fpage>&#x2013;<lpage>45</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/s11104-014-2229-z</pub-id>, PMID: <pub-id pub-id-type="pmid">41758449</pub-id>
</mixed-citation>
</ref>
<ref id="B20">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Loffredo</surname> <given-names>N.</given-names></name>
<name><surname>Mounier</surname> <given-names>S.</given-names></name>
<name><surname>Thiry</surname> <given-names>Y.</given-names></name>
<name><surname>Coppin</surname> <given-names>F.</given-names></name>
</person-group> (<year>2011</year>). 
<article-title>Sorption of selenate on soils and pure phases: kinetic parameters and stabilization</article-title>. <source>J. Environ. Radioactivity</source> <volume>102</volume>, <fpage>843</fpage>&#x2013;<lpage>851</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.jenvrad.2011.05.004</pub-id>, PMID: <pub-id pub-id-type="pmid">21683486</pub-id>
</mixed-citation>
</ref>
<ref id="B21">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Lu</surname> <given-names>D.</given-names></name>
<name><surname>Huang</surname> <given-names>T.</given-names></name>
<name><surname>Chen</surname> <given-names>J.</given-names></name>
<name><surname>Liao</surname> <given-names>Q.</given-names></name>
<name><surname>Wei</surname> <given-names>Y.</given-names></name>
<name><surname>Xing</surname> <given-names>Y.</given-names></name>
<etal/>
</person-group>. (<year>2023</year>). 
<article-title>Effects of biochar application on bioavailability in selenium-rich red soil areas</article-title>. <source>Chin. J. Soil Fertilizer</source> <volume>10)</volume>, <fpage>118</fpage>&#x2013;<lpage>126</lpage>.
</mixed-citation>
</ref>
<ref id="B22">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Lyu</surname> <given-names>C. H.</given-names></name>
<name><surname>Qin</surname> <given-names>Y. J.</given-names></name>
<name><surname>Zhao</surname> <given-names>Z. Q.</given-names></name>
<name><surname>Liu</surname> <given-names>X. W.</given-names></name>
</person-group> (<year>2021</year>). 
<article-title>Characteristics of selenium enrichment and assessment of selenium bioavailability using the diffusive gradients in thin-films technique in seleniferous soils in Enshi, Central China</article-title>. <source>Environ. pollut.</source> <volume>273</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.envpol.2021.116507</pub-id>, PMID: <pub-id pub-id-type="pmid">33493758</pub-id>
</mixed-citation>
</ref>
<ref id="B23">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Ma</surname> <given-names>S.</given-names></name>
<name><surname>Zhu</surname> <given-names>G.</given-names></name>
<name><surname>Parhat</surname> <given-names>R.</given-names></name>
<name><surname>Jin</surname> <given-names>Y.</given-names></name>
<name><surname>Wang</surname> <given-names>X.</given-names></name>
<name><surname>Wu</surname> <given-names>W.</given-names></name>
<etal/>
</person-group>. (<year>2023</year>). 
<article-title>Exogenous selenium and biochar application modulate the growth and selenium uptake of medicinal legume Astragalus species</article-title>. <source>Plants</source> <volume>12</volume>, <fpage>1957</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.3390/plants12101957</pub-id>, PMID: <pub-id pub-id-type="pmid">37653874</pub-id>
</mixed-citation>
</ref>
<ref id="B24">
<mixed-citation publication-type="book">
<person-group person-group-type="author">
<name><surname>Pilbeam</surname> <given-names>D. J.</given-names></name>
<name><surname>Greathead</surname> <given-names>H. M. R.</given-names></name>
<name><surname>Drihem</surname> <given-names>K.</given-names></name>
</person-group> (<year>2015</year>). &#x201c;
<article-title>Selenium</article-title>,&#x201d; in <source>A handbook of plant nutrition</source>, <edition>2nd edn</edition>. Eds. 
<person-group person-group-type="editor">
<name><surname>Barker</surname> <given-names>A. V.</given-names></name>
<name><surname>Pilbeam</surname> <given-names>D. J.</given-names></name>
</person-group> (
<publisher-name>CRC Press</publisher-name>, <publisher-loc>Boca Raton, FL</publisher-loc>), <fpage>165</fpage>&#x2013;<lpage>198</lpage>.
</mixed-citation>
</ref>
<ref id="B25">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Pu</surname> <given-names>S.</given-names></name>
<name><surname>Qian</surname> <given-names>X.</given-names></name>
<name><surname>Mao</surname> <given-names>X.</given-names></name>
<name><surname>Wang</surname> <given-names>L.</given-names></name>
<name><surname>He</surname> <given-names>Y.</given-names></name>
<name><surname>Qiu</surname> <given-names>G.</given-names></name>
<etal/>
</person-group>. (<year>2023</year>). 
<article-title>Study on the enrichment and translocation characteristics of selenium in crops in typical selenium-rich areas of Guizhou Province</article-title>. <source>Earth and Environment</source> <volume>51</volume>, <fpage>612</fpage>&#x2013;<lpage>619</lpage>.
</mixed-citation>
</ref>
<ref id="B26">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Qin</surname> <given-names>H.</given-names></name>
<name><surname>Jiang</surname> <given-names>D.</given-names></name>
<name><surname>Huang</surname> <given-names>X.</given-names></name>
<name><surname>Dang</surname> <given-names>H.</given-names></name>
<name><surname>Huang</surname> <given-names>J.</given-names></name>
<name><surname>Wang</surname> <given-names>M.</given-names></name>
</person-group> (<year>2023</year>). 
<article-title>Effects of organic matter on the adsorption and desorption characteristics of Se(IV) in acidic selenium-rich soil in Guangxi</article-title>. <source>Soil</source> <volume>55</volume>, <fpage>363</fpage>&#x2013;<lpage>371</lpage>.
</mixed-citation>
</ref>
<ref id="B27">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Qu</surname> <given-names>J.</given-names></name>
<name><surname>Xu</surname> <given-names>B.</given-names></name>
<name><surname>Gong</surname> <given-names>S.</given-names></name>
</person-group> (<year>1997</year>). 
<article-title>Determination of selenium speciation in soil and sediments by continuous extraction technique</article-title>. <source>Environ. Chem.</source> <volume>16</volume>, <fpage>277</fpage>&#x2013;<lpage>183</lpage>.
</mixed-citation>
</ref>
<ref id="B28">
<mixed-citation publication-type="book">
<person-group person-group-type="author">
<name><surname>Ran</surname> <given-names>H.</given-names></name>
</person-group> (<year>2023</year>). <source>Mechanism of arsenic sequestration and release in soils of metal mining areas by the transformation of iron and aluminum minerals</source> (<publisher-loc>Changsha</publisher-loc>: 
<publisher-name>Central South University</publisher-name>).
</mixed-citation>
</ref>
<ref id="B29">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Rayman</surname> <given-names>M. P.</given-names></name>
</person-group> (<year>2008</year>). 
<article-title>Food-chain selenium and human health: emphasis on intake</article-title>. <source>Br. J. Nutr.</source> <volume>100</volume>, <fpage>254</fpage>&#x2013;<lpage>268</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1017/S0007114508939830</pub-id>, PMID: <pub-id pub-id-type="pmid">18346308</pub-id>
</mixed-citation>
</ref>
<ref id="B30">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Sharma</surname> <given-names>S.</given-names></name>
<name><surname>Bansal</surname> <given-names>A.</given-names></name>
<name><surname>Dogra</surname> <given-names>R.</given-names></name>
<name><surname>Dhillon</surname> <given-names>S. K.</given-names></name>
<name><surname>Dhillon</surname> <given-names>K. S.</given-names></name>
</person-group> (<year>2011</year>). 
<article-title>Effect of organic amendments on uptake of selenium and biochemical grain composition of wheat and rape grown on seleniferous soils in northwestern India</article-title>. <source>J. Plant Nutr. Soil Sci.</source> <volume>174</volume>, <fpage>269</fpage>&#x2013;<lpage>275</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1002/jpln.200900265</pub-id>, PMID: <pub-id pub-id-type="pmid">41757603</pub-id>
</mixed-citation>
</ref>
<ref id="B31">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Stroud</surname> <given-names>J. L.</given-names></name>
<name><surname>Li</surname> <given-names>H. F.</given-names></name>
<name><surname>Lopez-Bellido</surname> <given-names>F. J.</given-names></name>
<name><surname>Broadley</surname> <given-names>M. R.</given-names></name>
<name><surname>Foot</surname> <given-names>I.</given-names></name>
<name><surname>Fairweather-Tait</surname> <given-names>S. J.</given-names></name>
<etal/>
</person-group>. (<year>2010</year>). 
<article-title>Impact of sulphur fertilisation on crop response to selenium fertilisation</article-title>. <source>Plant Soil</source> <volume>332</volume>, <fpage>31</fpage>&#x2013;<lpage>40</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/s11104-009-0230-8</pub-id>, PMID: <pub-id pub-id-type="pmid">41758449</pub-id>
</mixed-citation>
</ref>
<ref id="B32">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Tan</surname> <given-names>Z.</given-names></name>
<name><surname>Du</surname> <given-names>J.</given-names></name>
<name><surname>Sun</surname> <given-names>X.</given-names></name>
<name><surname>Yi</surname> <given-names>Q.</given-names></name>
<name><surname>Xu</surname> <given-names>P.</given-names></name>
<name><surname>Zhang</surname> <given-names>M.</given-names></name>
</person-group> (<year>2024</year>). 
<article-title>Effects of lime, phosphate and silicate on soil selenium availability and selenium accumulation in rice</article-title>. <source>Jiangsu J. Agric. Sci.</source> <volume>40</volume>, <fpage>450</fpage>&#x2013;<lpage>456</lpage>.
</mixed-citation>
</ref>
<ref id="B33">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Wang</surname> <given-names>D.</given-names></name>
<name><surname>Dinh</surname> <given-names>Q. T.</given-names></name>
<name><surname>Thu</surname> <given-names>T. T.</given-names></name>
<name><surname>Zhou</surname> <given-names>F.</given-names></name>
<name><surname>Yang</surname> <given-names>W.</given-names></name>
<name><surname>Wang</surname> <given-names>M.</given-names></name>
<etal/>
</person-group>. (<year>2018</year>). 
<article-title>Effect of selenium-enriched organic material amendment on selenium fraction transformation and bioavailability in soil</article-title>. <source>Chemosphere</source> <volume>199</volume>, <fpage>417</fpage>&#x2013;<lpage>426</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.chemosphere.2018.02.007</pub-id>, PMID: <pub-id pub-id-type="pmid">29453068</pub-id>
</mixed-citation>
</ref>
<ref id="B34">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Wang</surname> <given-names>D.</given-names></name>
<name><surname>Peng</surname> <given-names>Q.</given-names></name>
<name><surname>Yang</surname> <given-names>W. X.</given-names></name>
<name><surname>Dinh</surname> <given-names>Q. T.</given-names></name>
<name><surname>Tran</surname> <given-names>T. A. T.</given-names></name>
<name><surname>Zhao</surname> <given-names>X. D.</given-names></name>
<etal/>
</person-group>. (<year>2020</year>). 
<article-title>DOM derivations determine the distribution and bioavailability of DOM-Se in selenate applied soil and mechanisms</article-title>. <source>Environ. pollut.</source> <volume>259</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.envpol.2019.113899</pub-id>, PMID: <pub-id pub-id-type="pmid">31927276</pub-id>
</mixed-citation>
</ref>
<ref id="B35">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>White</surname> <given-names>P. J.</given-names></name>
</person-group> (<year>2016</year>). 
<article-title>Selenium accumulation by plants</article-title>. <source>Ann. Bot.</source> <volume>227</volume>, <fpage>217</fpage>&#x2013;<lpage>235</lpage>.
</mixed-citation>
</ref>
<ref id="B36">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>White</surname> <given-names>P. J.</given-names></name>
</person-group> (<year>2018</year>). 
<article-title>Selenium metabolism in plant</article-title>. <source>Biochim. Biophys. Acta Gen. Subj.</source> <volume>1862</volume>, <fpage>2333</fpage>&#x2013;<lpage>2342</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.bbagen.2018.05.006</pub-id>, PMID: <pub-id pub-id-type="pmid">29751098</pub-id>
</mixed-citation>
</ref>
<ref id="B37">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Winkel</surname> <given-names>L. H. E.</given-names></name>
<name><surname>Vriens</surname> <given-names>B.</given-names></name>
<name><surname>Jones</surname> <given-names>G. D.</given-names></name>
<name><surname>Schneider</surname> <given-names>L. S.</given-names></name>
<name><surname>Pilon-Smits</surname> <given-names>E.</given-names></name>
<name><surname>Ba&#xf1;uelos</surname> <given-names>G. S.</given-names></name>
</person-group> (<year>2015</year>). 
<article-title>Selenium cycling across soil-plant-atmosphere interfaces</article-title>. <source>A Crit. Rev. Nutrients</source> <volume>7</volume>, <fpage>4199</fpage>&#x2013;<lpage>4239</lpage>.
</mixed-citation>
</ref>
<ref id="B38">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Wu</surname> <given-names>Z.</given-names></name>
<name><surname>Li</surname> <given-names>M.</given-names></name>
<name><surname>Liang</surname> <given-names>H.</given-names></name>
<name><surname>Xing</surname> <given-names>R.</given-names></name>
<name><surname>Li</surname> <given-names>Y.</given-names></name>
<name><surname>Liu</surname> <given-names>C.</given-names></name>
</person-group> (<year>2025</year>). 
<article-title>Study on selenium enrichment characteristics of crops and soil-crop selenium correlation in selenium-rich areas of Anhui Province</article-title>. <source>Shandong Land Resour.</source> <volume>41</volume>, <fpage>31</fpage>&#x2013;<lpage>38</lpage>.
</mixed-citation>
</ref>
<ref id="B39">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Xie</surname> <given-names>B.</given-names></name>
<name><surname>He</surname> <given-names>L.</given-names></name>
<name><surname>Jiang</surname> <given-names>G.</given-names></name>
<name><surname>Sun</surname> <given-names>B.</given-names></name>
<name><surname>Zeng</surname> <given-names>D.</given-names></name>
<name><surname>Zhou</surname> <given-names>G.</given-names></name>
</person-group> (<year>2017</year>). 
<article-title>Regulation and evaluation of soil selenium availability in typical selenium-rich areas of southern China</article-title>. <source>Rock Mineral Anal.</source> <volume>36</volume>, <fpage>273</fpage>&#x2013;<lpage>281</lpage>.
</mixed-citation>
</ref>
<ref id="B40">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Yeasmin</surname> <given-names>M.</given-names></name>
<name><surname>Lamb</surname> <given-names>D.</given-names></name>
<name><surname>Choppala</surname> <given-names>G.</given-names></name>
<name><surname>Rahman</surname> <given-names>M. M.</given-names></name>
</person-group> (<year>2022</year>). 
<article-title>Impact of Sulfur on Biofortification and speciation of selenium in wheat grain grown in selenium-deficient soils</article-title>. <source>J. Soil Sci. Plant Nutr.</source> <volume>22</volume>, <fpage>3243</fpage>&#x2013;<lpage>3253</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/s42729-022-00882-0</pub-id>, PMID: <pub-id pub-id-type="pmid">41758449</pub-id>
</mixed-citation>
</ref>
<ref id="B41">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Zhang</surname> <given-names>D.</given-names></name>
<name><surname>Dong</surname> <given-names>T. Y.</given-names></name>
<name><surname>Ye</surname> <given-names>J.</given-names></name>
<name><surname>Hou</surname> <given-names>Z.</given-names></name>
</person-group> (<year>2017</year>). 
<article-title>Selenium accumulation in wheat (Triticum aestivum L) as affected by coapplication of either selenite or selenate with phosphorus</article-title>. <source>Soil Sci. Plant Nutr.</source> <volume>63</volume>, <fpage>37</fpage>&#x2013;<lpage>44</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1080/00380768.2017.1280377</pub-id>, PMID: <pub-id pub-id-type="pmid">41735180</pub-id>
</mixed-citation>
</ref>
<ref id="B42">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Zhang</surname> <given-names>L.</given-names></name>
<name><surname>Hu</surname> <given-names>B.</given-names></name>
<name><surname>Li</surname> <given-names>W.</given-names></name>
<name><surname>Che</surname> <given-names>R.</given-names></name>
<name><surname>Deng</surname> <given-names>K.</given-names></name>
<name><surname>Li</surname> <given-names>H.</given-names></name>
<etal/>
</person-group>. (<year>2014</year>). 
<article-title>OsPT2, a phosphate transporter, is involved in the active uptake of selenite in rice</article-title>. <source>New Phyto.</source> <volume>201</volume>, <fpage>1183</fpage>&#x2013;<lpage>1191</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/nph.12596</pub-id>, PMID: <pub-id pub-id-type="pmid">24491113</pub-id>
</mixed-citation>
</ref>
<ref id="B43">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Zhao</surname> <given-names>X.</given-names></name>
<name><surname>Lu</surname> <given-names>Y.</given-names></name>
<name><surname>Dai</surname> <given-names>L.</given-names></name>
<name><surname>Wang</surname> <given-names>L.</given-names></name>
<name><surname>Zhou</surname> <given-names>G.</given-names></name>
<name><surname>Liang</surname> <given-names>T.</given-names></name>
</person-group> (<year>2024</year>). 
<article-title>Selenium spatial distribution and bioavailability of soil-plant systems in China: A comprehensive review</article-title>. <source>Environ. Geochem Health</source> <volume>46</volume>, <fpage>341</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/s10653-024-02126-9</pub-id>, PMID: <pub-id pub-id-type="pmid">39073467</pub-id>
</mixed-citation>
</ref>
</ref-list>
<fn-group>
<fn id="n1" fn-type="custom" custom-type="edited-by">
<p>Edited by: <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/1927329">Mohammad Tarique Zeyad</ext-link>, National Bureau of Agriculturally Important Microorganisms (ICAR), India</p></fn>
<fn id="n2" fn-type="custom" custom-type="reviewed-by">
<p>Reviewed by: <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/2133037">Lei Yu</ext-link>, Yangzhou University, China</p>
<p><ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/3341225">Zebin Tan</ext-link>, Nanjing Agricultural University, China</p>
<p><ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/3348396">Mouloud Ait Mechedal</ext-link>, Universite Mohamed El Bachir El Ibrahimi de Bordj Bou Arreridj, Algeria</p></fn>
</fn-group>
</back>
</article>