<?xml version="1.0" encoding="utf-8"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD JATS (Z39.96) Journal Publishing DTD v1.3 20210610//EN" "JATS-journalpublishing1-3-mathml3.dtd">
<article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xmlns:ali="http://www.niso.org/schemas/ali/1.0/" article-type="research-article" dtd-version="1.3" xml:lang="EN">
<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Microbiol.</journal-id>
<journal-title-group>
<journal-title>Frontiers in Microbiology</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Microbiol.</abbrev-journal-title>
</journal-title-group>
<issn pub-type="epub">1664-302X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fmicb.2025.1734062</article-id>
<article-version article-version-type="Version of Record" vocab="NISO-RP-8-2008"/>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Original Research</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>Effects of different mulching practices on soil properties and soil microbial communities in tomato production</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Li</surname>
<given-names>Xiaoxia</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; review &#x0026; editing" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-review-editing/">Writing &#x2013; review &#x0026; editing</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Formal analysis" vocab-term-identifier="https://credit.niso.org/contributor-roles/formal-analysis/">Formal analysis</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="supervision" vocab-term-identifier="https://credit.niso.org/contributor-roles/supervision/">Supervision</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; original draft" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-original-draft/">Writing &#x2013; original draft</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="conceptualization" vocab-term-identifier="https://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="methodology" vocab-term-identifier="https://credit.niso.org/contributor-roles/methodology/">Methodology</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="visualization" vocab-term-identifier="https://credit.niso.org/contributor-roles/visualization/">Visualization</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="investigation" vocab-term-identifier="https://credit.niso.org/contributor-roles/investigation/">Investigation</role>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Cao</surname>
<given-names>Jinjun</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; original draft" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-original-draft/">Writing &#x2013; original draft</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="investigation" vocab-term-identifier="https://credit.niso.org/contributor-roles/investigation/">Investigation</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Formal analysis" vocab-term-identifier="https://credit.niso.org/contributor-roles/formal-analysis/">Formal analysis</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Data curation" vocab-term-identifier="https://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Li</surname>
<given-names>Dan</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="methodology" vocab-term-identifier="https://credit.niso.org/contributor-roles/methodology/">Methodology</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; original draft" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-original-draft/">Writing &#x2013; original draft</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="visualization" vocab-term-identifier="https://credit.niso.org/contributor-roles/visualization/">Visualization</role>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Jin</surname>
<given-names>Kunpeng</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="investigation" vocab-term-identifier="https://credit.niso.org/contributor-roles/investigation/">Investigation</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; original draft" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-original-draft/">Writing &#x2013; original draft</role>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Liu</surname>
<given-names>Yongzhong</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="methodology" vocab-term-identifier="https://credit.niso.org/contributor-roles/methodology/">Methodology</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; original draft" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-original-draft/">Writing &#x2013; original draft</role>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Li</surname>
<given-names>Wanxing</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>&#x002A;</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/3257539"/>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Formal analysis" vocab-term-identifier="https://credit.niso.org/contributor-roles/formal-analysis/">Formal analysis</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; review &#x0026; editing" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-review-editing/">Writing &#x2013; review &#x0026; editing</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="investigation" vocab-term-identifier="https://credit.niso.org/contributor-roles/investigation/">Investigation</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="conceptualization" vocab-term-identifier="https://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
</contrib>
</contrib-group>
<aff id="aff1"><label>1</label><institution>Millet Research Institute, Shanxi Agricultural University</institution>, <city>Changzhi</city>, <country country="cn">China</country></aff>
<aff id="aff2"><label>2</label><institution>Key Laboratory of Sustainable Dryland Agriculture of Shanxi Province</institution>, <city>Taiyuan</city>, <country country="cn">China</country></aff>
<author-notes>
<corresp id="c001"><label>&#x002A;</label>Correspondence: Wanxing Li, <email xlink:href="mailto:sxndjzs@126.com">sxndjzs@126.com</email></corresp>
</author-notes>
<pub-date publication-format="electronic" date-type="pub" iso-8601-date="2026-02-03">
<day>03</day>
<month>02</month>
<year>2026</year>
</pub-date>
<pub-date publication-format="electronic" date-type="collection">
<year>2025</year>
</pub-date>
<volume>16</volume>
<elocation-id>1734062</elocation-id>
<history>
<date date-type="received">
<day>30</day>
<month>10</month>
<year>2025</year>
</date>
<date date-type="rev-recd">
<day>29</day>
<month>12</month>
<year>2025</year>
</date>
<date date-type="accepted">
<day>30</day>
<month>12</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x00A9; 2026 Li, Cao, Li, Jin, Liu and Li.</copyright-statement>
<copyright-year>2026</copyright-year>
<copyright-holder>Li, Cao, Li, Jin, Liu and Li</copyright-holder>
<license>
<ali:license_ref start_date="2026-02-03">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>
<sec>
<title>Introduction</title>
<p>Agricultural mulches are commonly used for their benefits, however, the mechanisms by which they affect microbial communities to mediate soil properties that influence tomato growth remain unclear.</p>
</sec>
<sec>
<title>Methods</title>
<p>A three-year experiment was conducted comparing four treatments: plastic film mulching alone (SBF), straw mulching alone (SM), film-straw dual mulching (SBFSF), and no mulching (CK). Their effects on soil properties, microbial communities, and tomato growth were systematically evaluated.</p>
</sec>
<sec>
<title>Results</title>
<p>All mulching treatments significantly increased tomato yield, with SBF, SM, and SBFSF demonstrating improvement of 32.87%, 22.17%, and 50.17%, respectively. SBFSF exhibited the greatest dry matter weight (shoot plus root), root length, and root surface area 40 days post-transplanting, and it showed the strongest effect on soil moisture and thermoregulation. SM and SBFSF significantly increased soil organic carbon (SOC: +5.92%, +4.04%), total nitrogen (TN: +6.34%, +4.46%), available potassium (AK: +18.15%, +10.91%), and available phosphorus (AP: +2.60%, +2.15%). SBFSF significantly reduced the diversity of soil bacteria and fungi, howerer, it selectively increased the relative abundances of plant growth-promoting rhizobacteria and functional microorganisms involved in carbon-nitrogen cycling, such as the bacterial phylum Firmicutes(+125.18% to +193.55%), genera <italic>Lysobacter</italic> (+49.19% to +186.62%) and <italic>Bacillus</italic> (+168.56% to +273.00%), and fungal phyla <italic>Ascomycota</italic> (+7.99% to +8.19%) and <italic>Mortierellomycota</italic> (+11.05% to +98.71%), including the genus <italic>Trichocladium</italic> (+30.43% to +269.95%). In contrast, SM and SBF led to an increase in the abundance of pathogenic fungi (<italic>Fusarium</italic>, <italic>Cladosporium</italic>, <italic>Alternaria</italic>, and <italic>Cephaliophora</italic>), elucidating their inferior yield performance compared to SBFSF.</p>
</sec>
<sec>
<title>Discussion</title>
<p>Partial least squares path modeling (PLS-PM) analysis indicated that mulching practices directly and positively influenced the soil bacterial and fungal community composition and negatively affected soil fungal community diversity, which indirectly effecting tomato growth by modulating soil properties. These results provide a scientific foundation for improving mulching, and sustainable agricultural practices.</p>
</sec>
</abstract>
<kwd-group>
<kwd>mulching</kwd>
<kwd>soil bacteria</kwd>
<kwd>soil fungi</kwd>
<kwd>soil properties</kwd>
<kwd>tomato yield</kwd>
</kwd-group>
<funding-group>
<funding-statement>The author(s) declared that financial support was received for this work and/or its publication. This research was supported by the National Key Research and Development Program of China (No. 2021YFD1901101-5) and the project of Shanxi Province key lab construction (Z135050009017-1-10).</funding-statement>
</funding-group>
<counts>
<fig-count count="7"/>
<table-count count="1"/>
<equation-count count="0"/>
<ref-count count="58"/>
<page-count count="14"/>
<word-count count="9516"/>
</counts>
<custom-meta-group>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Terrestrial Microbiology</meta-value>
</custom-meta>
</custom-meta-group>
</article-meta>
</front>
<body>
<sec sec-type="intro" id="sec1">
<label>1</label>
<title>Introduction</title>
<p>Mulching practices play a crucial role in tomato cultivation for regulating soil hydrothermal conditions and the soil micro-environment (<xref ref-type="bibr" rid="ref5">Chakraborty et al., 2008</xref>; <xref ref-type="bibr" rid="ref46">Wr&#x00F3;blewska et al., 2019</xref>). The predominant mulching methods include plastic film mulching, straw mulching, net mulching with varying aeration, and biodegradable mulching films made from different materials. Plastic film mulching, net mulching, and biodegradable mulching films modify the surface energy balance and hydrothermal exchange, thereby promoting crop growth (<xref ref-type="bibr" rid="ref53">Zhao S. Q. et al., 2023</xref>). Conversely, Straw mulching (SM) reduces the need for chemical fertilizers, increases soil organic matter, improves soil structure, and facilitates nutrient cycling (<xref ref-type="bibr" rid="ref41">Wang et al., 2023</xref>; <xref ref-type="bibr" rid="ref19">Hu et al., 2024</xref>; <xref ref-type="bibr" rid="ref11">Du, 2024</xref>). Although mulching benefits are well documented and several studies have examined links among mulching, soil properties, microbial communities, and crop growth (<xref ref-type="bibr" rid="ref26">Luo et al., 2025</xref>; <xref ref-type="bibr" rid="ref17">Han et al., 2024</xref>), most work has evaluated a single mulching type. Few studies have systematically assessed the synergistic effects of a double mulching system (plastic film combined with straw) on soil microorganisms, soil physical and chemical properties, and crop performance. In particular, under specific regional soil and climatic conditions, the mechanisms by which combined plastic-film and straw mulching alters the soil environment and crop growth through regulation of key microbial groups remain unclear.</p>
<p>Soil microorganisms play an important role in soil (<xref ref-type="bibr" rid="ref33">Philippot et al., 2023</xref>), and their community composition, diversity and function directly affect soil fertility and plant health. Previous studies have shown that various mulching techniques can affect soil microbial communities by regulating the soil microenvironment, such as soil temperature, moisture, aeration, and carbon and nitrogen cycling, thereby potentially change the structure, diversity and function of soil microbial communities (<xref ref-type="bibr" rid="ref9">Dang et al., 2025</xref>; <xref ref-type="bibr" rid="ref8">Dai et al., 2024</xref>). Studies have demonstrated that straw mulching can provide abundant carbon sources that promote the growth of bacterial and fungal biomass and the activity of beneficial functional bacteria involved in carbon-nitrogen cycling (<xref ref-type="bibr" rid="ref42">Wang et al., 2024</xref>). However, the application of plastic mulches may lead to niche differentiation within microbial communities by creating distinct microenvironments, potentially favoring the growth of certain pathogenic bacteria (<xref ref-type="bibr" rid="ref34">Qi et al., 2022</xref>). Moreover, film-straw dual mulching has been shown to increase the abundance of straw-degrading bacteria, elevate soil organic content, enhance nutrients availability, and consequently improve crop yields (<xref ref-type="bibr" rid="ref48">Yang et al., 2024</xref>). However, the impacts of different mulching practices on the soil microenvironment vary significantly. The extent to which this difference, in particular selectively impacts the abundance and activity of beneficial (for instance plant rhizosphere growth promotors) or pathogenic bacteria, and how rhizospheric microorganisms react on this additional disturbance as well interact in one coherent system with soil nutrient dynamics in order to ultimately control growth and yield of crops, is an intriguing scientific question that should be resolved by present studies. In particular, the integrated film-straw dual mulching (SBFSF) method, which theoretically combines the advantages of both individual practices, lacks systematic empirical research regarding its comprehensive effects on soil microbiota and properties, as well as the underlying mechanisms.</p>
<p>Tomato, a crucial vegetable crop globally, is commonly cultivated under greenhouses using mulching technology to enhance growth conditions (<xref ref-type="bibr" rid="ref15">Gu et al., 2020</xref>). This study, based on a three-year experiment, investigates the impact of different mulching practices on soil properties and soil microbiology in tomato production. The specific objectives were: (1) to compare the impacts of film-straw dual mulching (SBFSF), plastic film mulch alone (SBF), straw mulch alone (SM), and no mulch (CK) on tomato plant growth and yield; (2) to examine changes in soil physical and chemical properties, and microbial communities under different mulching practices; and (3) to elucidate the mechanisms through which soil bacteria and fungi drive soil factors to promote tomato growth. The findings of this study are expected to provide a solid scientific basis and practical guidance for optimizing mulching techniques in tomato cultivation, steering the soil micro-environment, and harnessing the functional potential of soil microbes, thereby contributing to green and sustainable agricultural development.</p>
</sec>
<sec sec-type="materials|methods" id="sec2">
<label>2</label>
<title>Materials and methods</title>
<sec id="sec3">
<label>2.1</label>
<title>Site description and experimental design</title>
<p>The experiment was carried out at the Jindong Organic Dryland Farming Experimental Demonstration Base of Shanxi Agricultural University in Dianshang Town, Huguan County, Changzhi City, Shanxi Province (E113&#x00B0;27&#x2032;, N36&#x00B0;03&#x2032;), within a steel-framed arched plastic greenhouse (90&#x202F;m long, 8&#x202F;m wide, 3.5&#x202F;m high at the ridge, and 1.8&#x202F;m high at both sides, with natural ventilation). This site has a warm temperate semi-humid continental monsoon climate, with an average annual temperature of 10&#x202F;&#x00B0;C and rainfall of 574.5&#x202F;mm. The experiment was conducted in a completely randomized design with four treatments: no mulching (CK), straw mulching (SM, whole straw was covered on the ridges, 650&#x202F;kg&#x00B7;ha<sup>&#x2212;1</sup>), plastic film mulching (SBF), and combined film-straw mulching (SBFSF, plastic film on ridges with straw in furrows, 650&#x202F;kg&#x00B7;ha<sup>&#x2212;1</sup>). Each treatment had three replications, and every plot with five beds and five furrows. Planting specifications included two rows per bed, 35&#x202F;cm inter-plant spacing, 40&#x202F;cm inter-row spacing, 60&#x202F;cm ridge width, 80&#x202F;cm furrow width, and 22 tomato plants per row. Drip irrigation lines were placed along the planting rows. Tomato seedlings were transplanted on May 22, 2024, and the vines were removed on September 15, 2024. Standard agronomic practices, such as irrigation, pruning, and pesticide application (<xref rid="SM1" ref-type="supplementary-material">Supplementary Table S1</xref>), were uniformly applied to all treatments. The tomato variety tested was &#x201C;<italic>Rirun No. 8</italic>,&#x201D; utilizing 800&#x202F;mm wide, 0.01&#x202F;mm thick silver-black plastic film and corn straw from the previous year&#x2019;s harvest at the site for mulching. The initial soil physicochemical indicators at 0&#x2013;20&#x202F;cm soil layer are provided in <xref ref-type="table" rid="tab1">Table 1</xref>.</p>
<table-wrap position="float" id="tab1">
<label>Table 1</label>
<caption>
<p>Key properties of the experimental soil at 0&#x2013;20&#x202F;cm.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="top">Soil layer</th>
<th align="center" valign="top">SOC</th>
<th align="center" valign="top">Total nitrogen</th>
<th align="center" valign="top">Available phosphorus</th>
<th align="center" valign="top">Available potassium</th>
<th align="center" valign="top">Bulk density</th>
<th align="center" valign="top" rowspan="2">pH</th>
</tr>
<tr>
<th align="left" valign="middle">(cm)</th>
<th align="center" valign="middle">(g&#x00B7;kg<sup>&#x2212;1</sup>)</th>
<th align="center" valign="middle">(g&#x00B7;kg<sup>&#x2212;1</sup>)</th>
<th align="center" valign="middle">(mg&#x00B7;kg<sup>&#x2212;1</sup>)</th>
<th align="center" valign="middle">(mg&#x00B7;kg<sup>&#x2212;1</sup>)</th>
<th align="center" valign="middle">(g&#x00B7;cm<sup>&#x2212;3</sup>)</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="middle">0&#x2013;20</td>
<td align="center" valign="middle">33.2</td>
<td align="center" valign="middle">1.4</td>
<td align="center" valign="middle">13.42</td>
<td align="center" valign="middle">133.42</td>
<td align="center" valign="middle">1.25</td>
<td align="char" valign="middle" char=".">8.20</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="sec4">
<label>2.2</label>
<title>Sampling and measurement</title>
<sec id="sec5">
<label>2.2.1</label>
<title>Soil sample collection</title>
<p>To investigate the underlying mechanism by which mulching practices influences soil fertility and soil microbial community structure to improve tomato plant growth over a 3-year period, soil samples were collected in early September 2024 before tomato uprooting at the end of the experiments. The rhizosphere soil from five tomato plants was shaken off and mixed to form composite soil samples for each treatment, then passed through a 2&#x202F;mm sieve to remove gravel and residual roots. The soil samples were divided into two portions: one for analyzing soil chemical characteristics and the other stored at &#x2212;80&#x202F;&#x00B0;C for microbial DNA extraction (see <xref ref-type="fig" rid="fig1">Figure 1</xref>).</p>
<fig position="float" id="fig1">
<label>Figure 1</label>
<caption>
<p>Schematic diagram showing the fields conducted with four treatments.</p>
</caption>
<graphic xlink:href="fmicb-16-1734062-g001.tif" mimetype="image" mime-subtype="tiff">
<alt-text content-type="machine-generated">Four diagrams show different layouts of crop rows, each measuring seven meters by eight meters. The diagrams are labeled CK, SM, SBF, and SBFSF, with varying patterns of dark and light shading representing different planting schemes. Below each diagram is a corresponding photograph of a greenhouse with support structures and rows of plants matching the layout above.</alt-text>
</graphic>
</fig>
</sec>
<sec id="sec6">
<label>2.2.2</label>
<title>Soil physical and chemical properties measurement</title>
<p>Soil moisture content was assessed every 20&#x202F;days (avoiding periods of drip irrigation) by sampling using a 1&#x202F;m soil drill along the tomato rows, extracting soil samples at 20&#x202F;cm depth intervals from the surface to 1&#x202F;m. The samples were then oven-dried at 105&#x202F;&#x00B0;C until a constant weight and finally calculated the average gravimetric soil moisture content (g water/g dry soil) for the composite 1&#x202F;m depth sample. Soil bulk density was determined at harvest using the ring knife method, and he cutting ring had an inner diameter of 70&#x202F;mm and a height of 52&#x202F;mm. Soil temperature was recorded every 2&#x202F;h with a USB temperature logger (Jingchuang RC-5, Jiangsu Jingchuang Electric Co., LTD.), buried at depths of 10&#x202F;cm and 20&#x202F;cm within the tomato planting rows.</p>
<p>Soil organic carbon (SOC) was determined by digestion with 0.8&#x202F;M K<sub>2</sub>Cr<sub>2</sub>O<sub>7</sub> and concentrated H<sub>2</sub>SO<sub>4</sub>, followed by titration of residual dichromate (titration was performed using o-phenanthroline as the indicator). Total nitrogen (TN) was measured using an elemental analyzer (Vario EL III, Elementar, Germany). Available potassium (AK) was extracted with 1&#x202F;M NH&#x2084;OAc (pH 7.0) and measured by flame photometry. Available phosphorus (AK) was extracted with 0.5&#x202F;M NaHCO&#x2083; (pH 8.5) and determined by the molybdenum blue method, then quantified via molybdenum-antimony anti-UV spectrophotometric colorimetry (<xref ref-type="bibr" rid="ref2">Bao, 2000</xref>). The soil pH value was measured by a calibrated pH meter (FE28-Standard, Mettler-Toledo, Switzerland, soil-to-water ratio was 1:2.5).</p>
</sec>
<sec id="sec7">
<label>2.2.3</label>
<title>Soil DNA extraction and high-throughput sequencing</title>
<p>In this study, total DNA of soil bacteria and fungi was isolated using the OMEGA Soil DNA Kit (D5635-02) (Omega Bio-Tek, Norcross, GA, USA). Subsequently, the V3-V4 hypervariable regions of the bacterial 16S rRNA gene and the fungal ITS1 region were PCR amplified. The amplification products were electrophoresed on a 2% agarose gel, and target DNA fragments were recovered using the Axygen DNA Gel Extraction Kit. The PCR products were quantified with the Quant-iT PicoGreen dsDNA Assay Kit on a Microplate Reader (BioTek, FLx800). Library preparation and high-throughput sequencing were conducted by Personal Biotechnology Co., Ltd. The raw reads were deposited in the National Genomics Data Center, China National Center for Bioinformation (CRA037497).</p>
</sec>
</sec>
<sec id="sec8">
<label>2.3</label>
<title>Determination of dry matter weight of tomato plants, root morphology, and yields</title>
<p>The dry weight and root morphology in the 0&#x2013;20&#x202F;cm soil layer were monitored every 20&#x202F;days post transplantation of the tomato plants. Following this, the tomato plants were brought back to laboratory, where their dry weights were quantified after 105&#x202F;&#x00B0;C of exposure for 30&#x202F;min, drying them in an oven at 80&#x202F;&#x00B0;C until the constant weight and weighing the samples (<xref ref-type="bibr" rid="ref44">Wang F. et al., 2021</xref>). The root system determination method according to the following procedure: a custom-designed root extractor (in a cube shape, 20&#x202F;cm&#x202F;&#x00D7;&#x202F;20&#x202F;cm&#x202F;&#x00D7;&#x202F;20&#x202F;cm, 3&#x202F;mm thick, with one side shaped like a knife to facilitate soil penetration) was inserted into the soil at the tomato root location and driven in using a hammer after removing the aboveground portion of the plant. Subsequently, the root-soil block was then excavated with a spade and trimmed neatly using a professional cutting tool to obtain a standardized sample. The excavated root system along with the soil was transported to the laboratory, then carefully washed off the soil adhering to the roots. The root system was then submerged in a glass tank filled with water, where the roots were carefully separated and scanned using an LA2400 root scanner (Canadian Reagent Instruments Company) to collect data on total root length and root surface area. Finally, the scanned roots were then oven-dried at 80&#x202F;&#x00B0;C to a constant weight for calculating the dry matter weight of the root biomass.</p>
<p>During the harvest seasons of 2022&#x2013;2024, tomatoes were harvested in batches from the two middle rows of each plot, and the yield was calculated on a per-hectare basis.</p>
</sec>
<sec id="sec9">
<label>2.4</label>
<title>Data processing and analysis methods</title>
<p>The data were organized using Microsoft Excel 2007 (Microsoft Corp., USA), and variance analysis was performed with SPSS 27.0 (SPSS Inc., Chicago, USA), employing Duncan&#x2019;s method for significance testing, including tomato yield, dry matter weight, root morphology, soil physical and chemical properties as well as soil bacterial and fungal diversity index (all data were subjected to tests for normality and homogeneity of variances, and for datasets that did not meet the assumptions of normal distribution or homoscedasticity, appropriate data transformations were applied). Principal coordinates analysis (PCoA) was conducted using the Bray-Curtis distance matrix algorithm to compare the taxonomic composition among treatments. Functional prediction of microbial communities was conducted using PICRUSt2 software. Redundancy analysis (RDA) based on a linear model was performed to assess the effects of soil physical and chemical properties on the structure of soil bacterial and fungal communities. Partial least squares path model (PLS-PM) was used to explore relationships among soil physical properties, soil chemical properties, soil bacteria, soil fungi and tomato plant. Both RDA and PLS -PM analyses were implemented using the vegan and plspm packages in R language, respectively. Figures were generated using Origin 2022.</p>
</sec>
</sec>
<sec sec-type="results" id="sec10">
<label>3</label>
<title>Results</title>
<sec id="sec11">
<label>3.1</label>
<title>Tomato yields, dry matter weight and root morphology</title>
<sec id="sec12">
<label>3.1.1</label>
<title>Tomato yields</title>
<p>The results showed significant yield increases in all mulching treatments (SM, SBF, SBFSF) over the control during 2022&#x2013;2024 (SM vs. CK: <italic>p</italic> =&#x202F;0.010; SBF vs. CK: <italic>p</italic> =&#x202F;0.029; SBFSF vs. CK: <italic>p</italic> &#x003C;&#x202F;0.005), with three-year average gains of 17.11%&#x202F;&#x00B1;&#x202F;3.12%, 24.14%&#x202F;&#x00B1;&#x202F;7.53%, and 35.36%&#x202F;&#x00B1;&#x202F;7.82%, respectively. The annual increases were 11.42% (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.001), 9.15% (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.01), and 23.62% (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.001) in 2022 (<xref rid="SM1" ref-type="supplementary-material">Supplementary Figure S1</xref>) 17.74% (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.001), 30.40% (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.001), and 32.30% (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.001) in 2023 (<xref rid="SM1" ref-type="supplementary-material">Supplementary Figure S1</xref>); and 22.17% (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.001), 32.87% (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.001), and 50.17% (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.001) in 2024 (<xref ref-type="fig" rid="fig2">Figure 2a</xref>).</p>
<fig position="float" id="fig2">
<label>Figure 2</label>
<caption>
<p>Tomato yield <bold>(a)</bold>, dry matter weight <bold>(b)</bold>, and root morphology <bold>(c,d)</bold> under different mulching practices.</p>
</caption>
<graphic xlink:href="fmicb-16-1734062-g002.tif" mimetype="image" mime-subtype="tiff">
<alt-text content-type="machine-generated">Four bar graphs labeled a to d show agricultural metrics for treatments CK, SM, SBF, and SBFSF. Graph a displays yield, with SBFSF highest and CK lowest. Graph b shows dry matter weight across days 20 to 110, increasing with SBFSF leading. Graph c illustrates root length, progressively rising, with SBF and SBFSF having the longest lengths by day 110. Graph d depicts root surface area, similarly increasing, with SBFSF having the largest area by day 110. Different letters indicate statistically significant differences.</alt-text>
</graphic>
</fig>
</sec>
<sec id="sec13">
<label>3.1.2</label>
<title>Dry matter weight of tomato plants</title>
<p>In all stages of growth, the dry matter weight of plants under each mulching treatment were significantly higher than that of the control. Specifically, SM, SBF, and SBFSF showed an average increase of 14.54% (SM vs. CK: <italic>p</italic>&#x202F;&#x003E;&#x202F;0.05), 19.19% (SBF vs. CK: <italic>p</italic>&#x202F;&#x003C;&#x202F;0.001), and 23.44% (SBFSF vs. CK: <italic>p</italic>&#x202F;&#x003C;&#x202F;0.001), respectively, compared to CK. At 20&#x202F;days post-transplanting, the dry matter weight under SBF was significantly greater than that under SM and SBFSF, with no significant difference between SM and SBFSF. By 40&#x202F;days post-transplanting, SBFSF exhibited the highest dry matter weight, with no significant difference between SM and SBF (<xref ref-type="fig" rid="fig2">Figure 2b</xref>).</p>
</sec>
<sec id="sec14">
<label>3.1.3</label>
<title>Root morphology</title>
<p>Throughout the growth period, all mulching treatments demonstrated superior root length and surface area compared to CK. On average, SM, SBF, and SBFSF showed increases in root length of 13.85% (SM vs. CK: <italic>p</italic>&#x202F;&#x003C;&#x202F;0.001), 24.18% (SBF vs. CK: <italic>p</italic>&#x202F;&#x003C;&#x202F;0.001), and 25.09% (SBFSF vs. CK: <italic>p</italic>&#x202F;&#x003C;&#x202F;0.001), respectively, and in root surface area of 14.53% (SM vs. CK: <italic>p</italic>&#x202F;&#x003C;&#x202F;0.001), 15.76% (SBF vs. CK: <italic>p</italic>&#x202F;&#x003C;&#x202F;0.001), and 23.51% (SBFSF vs. CK: <italic>p</italic>&#x202F;&#x003C;&#x202F;0.001), respectively, relative to the control. Specifically, at 20&#x202F;days post-transplanting, SBF exhibited significantly greater root length and surface area compared to SM and SBFSF. By 40&#x202F;days post-transplanting, SBFSF displayed the highest values for both root length and surface area, with no significant difference observed compared to SM by 60&#x202F;days (<xref ref-type="fig" rid="fig2">Figures 2c</xref>,<xref ref-type="fig" rid="fig2">d</xref>).</p>
</sec>
</sec>
<sec id="sec15">
<label>3.2</label>
<title>Soil physical properties</title>
<sec id="sec16">
<label>3.2.1</label>
<title>Soil water content</title>
<p>At 20, 60, and 110&#x202F;days post-transplanting, soil moisture content under the mulching treatments consistently exhibited the following order: SBFSF &#x003E; SBF&#x202F;&#x003E;&#x202F;SM&#x202F;&#x003E;&#x202F;CK. The three mulching treatments increased soil moisture by 4.89&#x2013;12.9%, 5.14&#x2013;9.04%, and 1.51&#x2013;7.16%, respectively, compared to CK. At 40&#x202F;days after transplanting, SBF showed the highest moisture content, though no significant difference was observed compared to SBFSF. At 80&#x202F;days after transplanting, there were no significant differences among the three mulching treatments, yet all maintained higher moisture levels than CK (<xref ref-type="fig" rid="fig3">Figure 3a</xref>).</p>
<fig position="float" id="fig3">
<label>Figure 3</label>
<caption>
<p>Soil water content, soil temperature, and soil bulk density under different mulching practices: <bold>(a)</bold> Soil water content; <bold>(b)</bold> soil temperature at 10&#x202F;cm; <bold>(c)</bold> soil temperature at 20&#x202F;cm; <bold>(d)</bold> soil bulk density.</p>
</caption>
<graphic xlink:href="fmicb-16-1734062-g003.tif" mimetype="image" mime-subtype="tiff">
<alt-text content-type="machine-generated">Four-panel chart displaying soil characteristics under different treatments. Panel (a) shows bar graphs of soil water content (%) across five time intervals (20D-110D) with CK, SM, SBF, and SBFSF treatments. Panel (b) presents soil temperature (&#x00B0;C) over time using line graphs for each treatment. Panel (c) depicts soil temperature (&#x00B0;C) trends similarly, with all treatments plotted over time. Panel (d) shows soil bulk density (g/cm&#x00B3;) as bar graphs for each treatment, indicating significant differences marked by letters.</alt-text>
</graphic>
</fig>
</sec>
<sec id="sec17">
<label>3.2.2</label>
<title>Soil temperature</title>
<p>Soil temperature rose at the beginning of tomato growth period, and then declined with the slower fluctuations under deeper soil layer (<xref ref-type="fig" rid="fig3">Figures 3b</xref>,<xref ref-type="fig" rid="fig3">c</xref>). All mulching treatments maintained higher soil temperatures compared to the control (CK), with temperature increases ranging from 0.01&#x2013;2.21&#x202F;&#x00B0;C at 10&#x202F;cm depth and 0.02&#x2013;2.05&#x202F;&#x00B0;C at 20&#x202F;cm depth. The thermal dynamics displayed distinct temporal patterns: during the first 80&#x202F;days after transplantation, soil temperatures followed the order: SBF&#x202F;&#x003E;&#x202F;SBFSF &#x003E; SM&#x202F;&#x003E;&#x202F;CK. From 80 to 110 days, the temperature pattern shifted to: SBFSF &#x003E; SM&#x202F;&#x003E;&#x202F;SBF&#x202F;&#x003E;&#x202F;CK. In contrast to SBF, both SBFSF and SM treatments demonstrated thermoregulatory effects, providing cooling during high-temperature periods while maintaining warmth during cooler periods. SBFSF had the least soil temperature fluctuation over the full growth period of tomato. Correlation analysis showed no relationship between soil moisture and temperature at 20, 40, 60, and 80&#x202F;days after tomato transplanting, but a strong relationship emerged at 110&#x202F;days after transplanting, with a correlation coefficient of 0.966 (<italic>p</italic> &#x003C;&#x202F;0.001).</p>
</sec>
<sec id="sec18">
<label>3.2.3</label>
<title>Soil bulk density</title>
<p>The soil bulk density decreased by 2.40% (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.01) in the SM treatment compared to the CK treatment, whereas the SBF and SBFSF treatments demonstrated notable increases of 3.20% (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.01) and 2.40% (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.01), respectively (<xref ref-type="fig" rid="fig3">Figure 3d</xref>).</p>
</sec>
</sec>
<sec id="sec19">
<label>3.3</label>
<title>Soil chemical properties</title>
<p>Soil chemical properties exhibited significant change under different mulching treatments after 3&#x202F;years of tomato cultivation (<xref ref-type="fig" rid="fig4">Figure 4</xref>). Compared to the control (CK), SM and SBFSF significantly increased the contents of soil organic matter, total nitrogen, available potassium, and available phosphorus, with corresponding increases of 5.92%, 6.34%, 18.15%, and 2.60% (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.01) for SM, and 4.04%, 4.46%, 10.91%, and 2.15% (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.01) for SBFSF. In contrast, SBF resulted in a significant reduction in soil organic matter content by 3.85% (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.01), though it increased available potassium content by 4.98% (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.01). All mulching treatments significantly reduced soil pH.</p>
<fig position="float" id="fig4">
<label>Figure 4</label>
<caption>
<p>Characteristics of soil chemical properties under different mulching practices.</p>
</caption>
<graphic xlink:href="fmicb-16-1734062-g004.tif" mimetype="image" mime-subtype="tiff">
<alt-text content-type="machine-generated">Bar charts display various soil properties under different treatments: CK, SM, SBF, and SBFSF. The properties measured are organic matter, total nitrogen, available potassium, available phosphorus, and pH. Each bar is differentiated by color and accompanied by statistical annotations.</alt-text>
</graphic>
</fig>
</sec>
<sec id="sec20">
<label>3.4</label>
<title>Microbial communities and potential metabolism functions</title>
<p>SBFSF significantly increased only the fungal Chao1 index, while showing significant reductions in all other alpha-diversity indices for both bacterial and fungal communities. In contrast, SM exhibited significant increases across all measured alpha-diversity indices. Regarding SBF, no significant difference was observed in the bacterial Chao 1 index compared to CK, however, all remaining alpha-diversity indices showed statistically significant enhancement (<xref ref-type="fig" rid="fig5">Figures 5a</xref>&#x2013;<xref ref-type="fig" rid="fig5">d</xref>).</p>
<fig position="float" id="fig5">
<label>Figure 5</label>
<caption>
<p>Soil bacterial and fungi diversity index (based on OTU level). <bold>(a)</bold> Chao 1 index of bacteria, <bold>(b)</bold> Shannon 1 index of bacteria, <bold>(c)</bold> Chao 1 index of fungi, <bold>(d)</bold> Shannon 1 index of fungi, <bold>(e)</bold> principal coordinate analysis (PCoA) of bacteria, <bold>(f)</bold> principal coordinate analysis (PCoA) of fungi.</p>
</caption>
<graphic xlink:href="fmicb-16-1734062-g005.tif" mimetype="image" mime-subtype="tiff">
<alt-text content-type="machine-generated">Six graphs showing data on groups labeled CK, SM, SBF, and SBFSF. The graphs (a) and (c) display Chao1 indices with different levels marked by letters. Graphs (b) and (d) show Shannon indices. Graphs (e) and (f) are PCoA plots illustrating differences among the groups. Each graph is color-coded: CK (red), SM (blue), SBF (green), and SBFSF (purple).</alt-text>
</graphic>
</fig>
<p>The composition of bacterial and fungal communities in soil was different under various mulching treatments, as shown by the Principal Coordinates Analysis (PCoA). The first axis of variation for the bacterial community explained 23.5%, while the second axis explained 21.4% in community structure. The values for all of the mulching treatments had a clear separation from control (CK) (<xref ref-type="fig" rid="fig5">Figure 5e</xref>). In contrast, although the fungal communities under various mulching treatments did not overlap with CK in the ordination space, the SBFSF treatment was relatively closer to CK. The first and second axes explained 59.7 and 20.4% of the variance in fungal community composition, respectively (<xref ref-type="fig" rid="fig5">Figure 5f</xref>).</p>
<p>The composition and the relative abundance of soil bacteria were analyzed at both the phylum (top 10) and genus (top 10) levels. Compared to CK, SBFSF, SM and SBF significantly reduced the relative abundance of <italic>Proteobacteria</italic> by &#x2212;17.20% to &#x2212;4.60%. Compared to other treatments, SBFSF significantly increased the relative abundance of <italic>Firmicutes</italic> by +125.18 to 193.55%, while SM enhanced <italic>Actinobacteria</italic> (+2.63% to +22.04%) and <italic>Chloroflexi</italic> (+50.70&#x2013;71.87%). SBF showed a significant increase in <italic>Acidobacteria</italic> (+21.79&#x2013;37.12%) (<xref ref-type="fig" rid="fig6">Figure 6a</xref>). At the genus level, SBFSF enriched <italic>Lysobacter</italic> (+49.19% to +186.62%) and <italic>Bacillus</italic> (+168.56% to +273.00%), whereas reduced the relative abundance of <italic>Sphingomonas</italic> (&#x2212;28.32% to &#x2212;9.60%). SM increased <italic>JG30-KF-CM45</italic> (+59.61% to +73.40%) and <italic>Arthrobacter</italic> (+28.94% to +55.42%), whereas reduced the relative abundance of <italic>Sphingomonas</italic> (&#x2212;46.51% to &#x2212;25.37%). SBF led to higher relative abundance of <italic>Vicinamibacteraceae</italic> (+19.34% to +41.39%) (<xref ref-type="fig" rid="fig6">Figure 6b</xref>).</p>
<fig position="float" id="fig6">
<label>Figure 6</label>
<caption>
<p>Relative abundance of bacterial communities at the phylum <bold>(a)</bold> and genus <bold>(b)</bold> levels, relative abundance of fungal communities at the phylum <bold>(c)</bold> and genus levels <bold>(d)</bold>, functional prediction and differences of bacterial <bold>(e)</bold> and fungal <bold>(f)</bold> communities under different mulching practices, correlation analyses between bacterial functions and genera <bold>(g)</bold>; correlation analyses between fungal functions and genera <bold>(h)</bold>. FSD, Folding, sorting and degradation; TPM, Metabolism of terpenoids and polyketides; OACM, Metabolism of other amino acids; CM, Carbohydrate metabolism; EM, Energy metabolism; CVM, Metabolism of cofactors and vitamins; RR, Replication and repair; XBM, Xenobiotics biodegradation and metabolism; LM, Xenobiotics biodegradation and metabolism; ACM, Amino acid metabolism; AAB, Amino acid biosynthesis; FALB, Fatty acid and lipid biosynthesis; Fer, Fermentation; CPEVB, Electron carrier, and vitamin biosynthesis; CB, Carbohydrate biosynthesis; SMB, Secondary metabolite biosynthesis; Res, Respiration; ET, Electron transfer; PPP, Pentose phosphate pathways; NNB, Nucleoside and nucleotide biosynthesis.</p>
</caption>
<graphic xlink:href="fmicb-16-1734062-g006.tif" mimetype="image" mime-subtype="tiff">
<alt-text content-type="machine-generated">The image contains multiple charts depicting microbial diversity and abundance across different treatments. Panels a) and b) are stacked bar graphs showing the relative abundance of bacterial phyla and genera. Panels c) and d) display similar information for fungal phyla and genera. Panels e) and f) are heat maps indicating the hierarchical clustering of sample treatments based on microbial composition. Panels g) and h) present correlation matrices for microbial relationships among the treatments. Each panel includes labels for treatments CK, SM, SBF, and SBFSF, with color codes indicating different microbial groups.</alt-text>
</graphic>
</fig>
<p>Fungal community analysis revealed that <italic>Ascomycota</italic> was the dominant phylum with relative abundances ranging from 86.76 to 94.38% (<xref ref-type="fig" rid="fig6">Figure 6c</xref>). Compared to other treatments, SBFSF significantly increased the relative abundances of <italic>Ascomycota</italic> (+7.99% to +8.19%) and <italic>Mortierellomycota</italic> (+11.05% to +98.71%). At the genus level, the relative abundance of <italic>Trichocladium</italic> in SBFSF was significantly higher than that in other treatments, showing an increase of 30.43 to 269.95%, while SM treatment notably enriched <italic>Fusarium</italic> (+120.39% to +149.16%), <italic>Cladosporium</italic> (587.22% to +4141.65%), and <italic>Alternaria</italic> (10953.15% to +32375.165). Moreover, SBF displayed significant increases in both <italic>Cephaliophora</italic> (+218.39% to +454.82%) and <italic>Fusarium</italic> (+12.48% to +180.25%) (<xref ref-type="fig" rid="fig6">Figure 6d</xref>).</p>
<p>Distinct soil microbial taxa enriched in various mulching treatments were identified through LEfSe analysis (<xref rid="SM1" ref-type="supplementary-material">Supplementary Figure S2</xref>). Overall, the numbers of indicator bacterial taxa associated decreased across all mulching types. Specifically, the SM and SBFSF management strategies promoted the enrichment of fungal taxa, while the SBF practice had a diminishing effect. Notably, the bacterial genus <italic>Lysobacter</italic> showed enrichment in response to SBFSF treatment, while <italic>Chloroflexi</italic> predominated under SM and <italic>Acidobacteriota</italic> under SBF. Among fungal taxa, <italic>Trichocladium</italic> was enriched in SBFSF, <italic>Dothideomycetes</italic> in SM, and <italic>Microascales</italic> in SBF.</p>
<p>Functional prediction of bacterial metabolism revealed distinct metabolic profiles in SBFSF compared to SM and SBF, with enhanced activities in amino acid metabolism, carbohydrate metabolism, metabolism of other amino acids, and lipid metabolism (<xref ref-type="fig" rid="fig6">Figure 6E</xref>). For fungal metabolic functions, SBFSF demonstrated increased capabilities in multiple pathways relative to SM and SBF, including electron transfer, respiration, secondary metabolite biosynthesis, cofactor and prosthetic group biosynthesis, electron carrier synthesis, vitamin biosynthesis, fermentation, as well as acid and lipid biosynthesis (<xref ref-type="fig" rid="fig6">Figure 6F</xref>).</p>
<p>At the bacterial genus level, <italic>Bacillus</italic> showed a significant positive correlation with metabolism of other amino acids, with a notably high correlation coefficient of 0.83 for carbohydrate metabolismz although not statistically significant. <italic>Lysobacter</italic> exhibited strong correlations with amino acid metabolism (0.85), lipid metabolism (0.76), and metabolism of other amino acids (0.86). Additionally, <italic>67&#x2013;14</italic> showed a significant positive correlation with carbohydrate metabolism and a notably high correlation coefficient of 0.70 for other amino acid metabolism (<xref ref-type="fig" rid="fig6">Figure 6G</xref>). At the fungal genus level, <italic>Mycochlamys</italic> demonstrated substantial correlations with electron carrier and vitamin biosynthesis (0.92), electron transfer (0.84), respiration (0.84), and carbohydrate biosynthesis (0.88) (<xref ref-type="fig" rid="fig6">Figure 6H</xref>).</p>
</sec>
<sec id="sec21">
<label>3.5</label>
<title>Relationship between microbial communities, soil properties, and tomato plant growth</title>
<p>Redundancy analysis conducted at the bacterial genus level, focusing on the top five scoring bacteria, revealed that the two principal axes explained 49.2 and 36.2% of the total variance contribution, respectively (<xref ref-type="fig" rid="fig7">Figure 7a</xref>). Specifically, the application of SBFSF was found to enhance the proliferation of <italic>Bacillus</italic> and <italic>Lysobacter</italic>. <italic>Bacillus</italic> exhibited a significant positive correlation with soil water content (SWC) (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.001) and a negative correlation with pH (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.05). Moreover, the use of SM was observed to stimulate the growth of <italic>JG30-KF-CM45</italic>, which displayed a negative correlation with soil bulk density (SBD) (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.05). Additionally, <italic>Sphingomonas</italic> showed negative correlations with available potassium (AK), available phosphorus (AP), total nitrogen (TN), and soil organic matter (SOM) (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.01). Similarly, redundancy analysis conducted at the fungal genus level, focusing on the top five fungi, in relation to soil environmental factors, indicated that the two principal axes explained 48.8 and 29.1% of the total variance, respectively (<xref ref-type="fig" rid="fig7">Figure 7b</xref>). The application of SBFSF was found to promote the growth of <italic>Trichocladium</italic>, while SM significantly enhanced the growth of <italic>Alternaria</italic>. <italic>Alternaria</italic> exhibited a positive correlation with total nitrogen (TN) (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.05) and a negative correlation with soil bulk density (SBD) (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.01). Furthermore, the use of SBF was associated with increased growth of <italic>Cephalophora</italic>, which displayed negative correlations with soil organic matter (SOM), total nitrogen (TN), and available potassium (AK) (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.05).</p>
<fig position="float" id="fig7">
<label>Figure 7</label>
<caption>
<p>RDA of soil bacterial <bold>(a)</bold> and fungi <bold>(b)</bold> communities in relation to environmental factors, partial least squares path model (PLS-PM) direct and indirect effects of bacteria community <bold>(c)</bold>, fungal community <bold>(d)</bold> and mulching practices on tomato yield. Solid lines denote positive correlations and dashed lines denote negative correlations. Numbers above lines are the standardized path coefficients. GOF, goodness-of-fit. &#x002A;, <italic>p</italic>&#x202F;&#x003C;&#x202F;0.050 level; &#x002A;&#x002A;, <italic>p</italic>&#x202F;&#x003C;&#x202F;0.010 level; &#x002A;&#x002A;&#x002A;, <italic>p</italic>&#x202F;&#x003C;&#x202F;0.001 level. Bacterial community compositions suggest the <italic>Proteobacteria</italic> and <italic>Actinobacteriota, Acidobacteriota, Chloroflexi, Bacteroidota</italic>, respectively. Fungal community compositions suggest the <italic>Ascomycota</italic> and <italic>Mortierellomycota</italic>. Community diversity depict the Chao 1 and Shannon indices of the bacteria and fungi. Soil properties suggest the soil organic matter, total nitrigon, available potassium, available phosphorus, pH, soil bulk density, soil temperature and soil water content, respectively. Plant growth suggest the dry matter weight, root length, root surface area, respectively.</p>
</caption>
<graphic xlink:href="fmicb-16-1734062-g007.tif" mimetype="image" mime-subtype="tiff">
<alt-text content-type="machine-generated">Two paired diagrams are presented. Panels (a) and (b) display redundancy analysis plots, showing relationships among different treatments (CK, SBF, SBFSF, SM) and variables such as pH, TN, and various bacteria. Panels (c) and (d) illustrate path analysis models of the effects of mulching practices on soil properties, community diversity, plant growth, and yield. Both models indicate significant relationships and explain variations, with different R-squared values for each component. Key pathways and their significance levels are highlighted with different line styles.</alt-text>
</graphic>
</fig>
<p>Further analysis using partial least squares path modeling (PLS-PM) revealed that a combination of soil bacterial community composition, diversity, soil properties, and plant growth accounted for 60.3% of the variability in tomato yield (<xref ref-type="fig" rid="fig7">Figure 7c</xref>). Similarly, soil fungal community composition, diversity, soil properties, and plant growth elucidated 60.8% of the variance in tomato yield (<xref ref-type="fig" rid="fig7">Figure 7d</xref>). Mulching practices exerted a direct and significant positive effect on soil bacterial community composition (path coefficient&#x202F;=&#x202F;0.578, <italic>p</italic>&#x202F;&#x003C;&#x202F;0.010) and diversity (path coefficient&#x202F;=&#x202F;0.599, <italic>p</italic>&#x202F;&#x003C;&#x202F;0.010), but only bacterial community composition had significant negative impacts on soil properties (path coefficient&#x202F;=&#x202F;&#x2212;0.804, <italic>p</italic>&#x202F;&#x003C;&#x202F;0.001), which subsequently positively influenced tomato plant growth (path coefficient&#x202F;=&#x202F;1.375, <italic>p</italic>&#x202F;&#x003C;&#x202F;0.001) and ultimately enhanced tomato yield (path coefficient&#x202F;=&#x202F;0.923, <italic>p</italic>&#x202F;&#x003C;&#x202F;0.001). The mulching practices exerted a direct and positive effect on the composition of the soil fungal community (path coefficient&#x202F;=&#x202F;0.806, <italic>p</italic>&#x202F;&#x003C;&#x202F;0.001), consequently positively influenced soil properties (path coefficient&#x202F;=&#x202F;0.406, <italic>p</italic>&#x202F;&#x003C;&#x202F;0.050). Conversely, it negatively affected fungal diversity (path coefficient&#x202F;=&#x202F;&#x2212;0.708, <italic>p</italic>&#x202F;&#x003C;&#x202F;0.001), which subsequently negatively affected soil properties (path coefficient&#x202F;=&#x202F;&#x2212;0.417, <italic>p</italic>&#x202F;&#x003C;&#x202F;0.050). The improved soil properties then positively promoted tomato plant growth (path coefficient&#x202F;=&#x202F;0.870, <italic>p</italic>&#x202F;&#x003C;&#x202F;0.001), ultimately benefiting tomato plant growth (path coefficient&#x202F;=&#x202F;0.924, <italic>p</italic>&#x202F;&#x003C;&#x202F;0.001).</p>
</sec>
</sec>
<sec sec-type="discussion" id="sec22">
<label>4</label>
<title>Discussion</title>
<sec id="sec23">
<label>4.1</label>
<title>Effect of different mulching practices on soil physicochemical properties</title>
<p>Previous studies has shown that both plastic film mulching and straw mulching are both able to prevent water movement and atmospheric gaseous exchange in soil (<xref ref-type="bibr" rid="ref4">Chai et al., 2022</xref>; <xref ref-type="bibr" rid="ref51">Zhang et al., 2021</xref>; <xref ref-type="bibr" rid="ref59">Zribi et al., 2015</xref>), thus enhancing soil hydrothermal properties compared with bare plots. Our study found that all three mulching treatments enhanced soil water retention capacity, with SBFSF exhibiting the highest moisture retention due to the complementary functions of its components within a fully covered microenvironment. At 80&#x202F;days after transplantation, no significant differences in soil water content were detected among the three mulching treatments. This result may be explained by the high transpiration rates of tomato plants under all mulching practices during this stage, leading to increased soil water depletion and consequently diminished variation in soil moisture levels.</p>
<p>Mulching practices enhanced soil temperature, with increases ranging between 0.01&#x2013;2.21&#x202F;&#x00B0;C at 10&#x202F;cm depth and 0.02&#x2013;2.05&#x202F;&#x00B0;C at 20&#x202F;cm depth, which is in agreement with former study results (<xref ref-type="bibr" rid="ref30">Miao et al., 2025</xref>; <xref ref-type="bibr" rid="ref45">Wang Y. et al., 2025</xref>; <xref ref-type="bibr" rid="ref35">Qu and Feng, 2022</xref>). In this study, both SBFSF and SM demonstrated the cooling effect under high temperatures and warming effect under low temperatures, indicating that the soil temperature regulation effects of different mulching practices exhibited temporal variations. SBF exhibited the strongest warming effect in the early stage of tomato growth, but it could not regulate temperature during high-temperature period. In contrast, SBFSF maintained the most stable soil thermal conditions during the entire growth period, which implied combined effect of full soil coverage on decreasing temperature fluctuation. These results emphasized the superior performance of SBFSF in generating a balanced and flexible thermal environment and such characteristics might contribute to enhancing tomato growth and yield stability under dynamic climate.</p>
<p>Our study demonstrated that both SM and SBFSF enhanced rhizosphere soil fertility after 3&#x202F;years of tomato cultivation by increasing the content of soil organic matter, total nitrogen, and available nutrients (<xref ref-type="bibr" rid="ref58">Zhu et al., 2022</xref>). This improvement occurred because straw mulching on the soil surface releases nutrients through microbial decomposition (<xref ref-type="bibr" rid="ref25">Liu et al., 2014</xref>), thereby enriching the topsoil nutrient pool&#x2014;a finding consistent with the research of <xref ref-type="bibr" rid="ref3">Biswal et al. (2022)</xref>, <xref ref-type="bibr" rid="ref43">Wang J. et al. (2021)</xref>, and <xref ref-type="bibr" rid="ref7">Chen et al. (2022)</xref>. SM treatment reduced the relative abundance of <italic>Sphingomonas</italic>, and RDA analysis indicated that <italic>Sphingomonas</italic> was negatively correlated with soil available potassium, available phosphorus, and organic matter (<italic>p</italic> &#x003C;&#x202F;0.01). Therefore, it can be inferred that <italic>Sphingomonas</italic> played an important role in the soil fertility of SM treatment. The improvement in soil fertility under SBFSF was also associated with its regulation on soil temperature. Small fluctuation of soil temperature in return leads to the low mineralization rates of SOC and available nutrients (<xref ref-type="bibr" rid="ref57">Zhou et al., 2012</xref>). We attribute the increased soil fertility under SBFSF to the combined regulatory effects of plastic film on ridges and straw in furrows. SBFSF likely produces a more favorable soil microenvironment which together support improved soil fertility. In addition, SM reduced soil bulk density which contributed to crop growth and roots residue input into soil for conversion into plant available nitrogen and phosphorous nutrient (<xref ref-type="bibr" rid="ref42">Wang et al., 2024</xref>). Under SBF, initially higher soil temperatures enhance the process of organic matter decomposition and exhaustion of available nutrient resources at a faster rate, leading to lower rhizosphere fertility.</p>
</sec>
<sec id="sec24">
<label>4.2</label>
<title>Effect of different mulching practices on soil microbial community diversity</title>
<p>Soil microorganisms are the most dynamic components of soil, with their community diversity and composition being influenced by mulching practices. In the present study, all mulching treatments significantly affected the richness and diversity of rhizosphere soil bacterial and fungal populations, and was due to the difference in soil temperature, moisture, and organic matter input by mulching practices (<xref ref-type="bibr" rid="ref27">Luo et al., 2019</xref>; <xref ref-type="bibr" rid="ref23">Ling et al., 2016</xref>; <xref ref-type="bibr" rid="ref6">Chen et al., 2017</xref>). SBFSF significantly increased fungal Chao1 index and decreased all other alpha-diversity indices for both bacteria and fungi. This was an indication that although SBFSF could promote the richness of fungus, it might also act as a limited ecological niche and inhibit overall microbial diversity (<xref ref-type="bibr" rid="ref52">Zhao et al., 2024</xref>). On the contrary, SM caused significant increases in all alpha-diversity measures which may indicate that a niche is less selective for richness and evenness. A majority of the previous reports suggested that straw mulching increased bacterial diversity (<xref ref-type="bibr" rid="ref49">Zhang et al., 2024</xref>), while results on fungal communities have not been consistent (<xref ref-type="bibr" rid="ref13">Fu et al., 2019</xref>; <xref ref-type="bibr" rid="ref24">Liu et al., 2024</xref>) due to variation in source and time of straw application and soil type, climate condition. Soil bacteria and fungi diversity were significantly increased by SBF, partially in line with <xref ref-type="bibr" rid="ref10">Dong et al. (2017)</xref>. The increase in soil bacterial and fungal diversity under the SBF may be attributed to the enhanced heterogeneity of the soil micro-environment induced by plastic film mulching. Variations in temperature, moisture, and oxygen availability across different soil layers create distinct niches, which can support the proliferation of microorganisms occupying diverse ecological roles (<xref ref-type="bibr" rid="ref21">Li et al., 2024</xref>).</p>
<p>In our study, SBFSF was effective in enhancing numbers of bacterial genera with biocontrol potential and nutrient cycling properties. The SBFSF enriched <italic>Bacillus</italic> and <italic>Lysobacter</italic>, which correlated with the soil carbon and nitrogen cycling. <italic>Bacillus</italic> are also well known and have been reported to possess a potential for antagonism against soil-borne pathogens, mediated by the production of antimicrobial compounds (<xref ref-type="bibr" rid="ref36">Saxena et al., 2020</xref>), this in turn could enhance crop productivity and carbon sequestration (<xref ref-type="bibr" rid="ref39">Wang et al., 2015</xref>). The predicted functional linkage to the metabolism of other amino acids strengthens the inference for their greater contribution to N cycling and the biosynthesis of bioactive compounds. Similarly, <italic>Lysobacter</italic> can also lyse fungi and are involved in the decomposition of resistant organic matter (<xref ref-type="bibr" rid="ref1">Afoshin et al., 2020</xref>). Their predicted functional profiles, associated with amino acid metabolism, lipid metabolism, and metabolism of other amino acids, suggested their multifunctional potential in C and N turnover. These findings suggested that SBFSF build a disease-suppressive soil environment with continuous nutrient cycling. In contrast, SM significantly increased <italic>Actinobacteriota</italic> and <italic>Chloroflexi</italic> but also enriched potentially pathogenic fungi. It is well known that members of <italic>Actinobacteriota</italic> can degrade complex lignocellulose and produce antibiotic compounds, which suppress the growth and reproduction of the pathogenic microorganisms (<xref ref-type="bibr" rid="ref48">Yang et al., 2024</xref>; <xref ref-type="bibr" rid="ref29">Ma et al., 2018</xref>; <xref ref-type="bibr" rid="ref56">Zheng et al., 2022</xref>), <italic>Chloroflexi</italic> involving organic substance decomposition, carbon cycling to increase soil conditions making it more suitable for plant growth (<xref ref-type="bibr" rid="ref18">Hu et al., 2023</xref>). At the genus level, SM promoted the substantial growth of <italic>JG30-KF-CM45</italic>, suggesting that <italic>JG30-KF-CM45</italic> may play an important role in soil carbon and nitrogen cycling. However, the substantial increase in <italic>Fusarium</italic>, <italic>Cladosporium</italic>, and <italic>Alternaria</italic> - species all containing known pathogens causing various disorders of tomato like wilt, leaf spot and fruit rot (<xref ref-type="bibr" rid="ref55">Zhao L. et al., 2023</xref>; <xref ref-type="bibr" rid="ref20">Kong et al., 2023</xref>; <xref ref-type="bibr" rid="ref50">Zhang et al., 2025</xref>)&#x2014;were alarming with respect to the possible risk potential of diseases due to SM. Thus, while the effect of SM on beneficial decomposers such as <italic>Actinobacteriota</italic> and <italic>Chloroflexi</italic> are promoted, the increase in plant potential pathogens appears to counterbalance those beneficial ecological effects. SBF significantly enriched <italic>Acidobacteria</italic> and <italic>Vicinamibacteraceae</italic>, taxa commonly associated with acidic environments (<xref ref-type="bibr" rid="ref14">Gon&#x00E7;alves et al., 2024</xref>), a finding consistent with the decreased soil pH observed under SBF in our study. Furthermore, SBF increased the relative abundance of <italic>Fusarium</italic> and <italic>Cephaliophora</italic>. Of particular note, a strain of <italic>Cephaliophora</italic> known to cause tomato leaf spot was first reported in the United States in 2023 (<xref ref-type="bibr" rid="ref28">Luo et al., 2020</xref>), suggesting that long-term silver-black plastic film mulching may also promote the proliferation of potential pathogenic fungi. It is noteworthy that the identification of the aforementioned pathogens is based on an analysis of relevant literature, and their precise functions require further experimental validation.</p>
<p>Fungal community analysis revealed that SBFSF increased the abundance of <italic>Ascomycota</italic>, <italic>Mortierellomycota</italic> at the phylum level and <italic>Trichocladium</italic> at the genus level, which are involved in decomposition of cellulose and chitin (<xref ref-type="bibr" rid="ref22">Li et al., 2020</xref>; <xref ref-type="bibr" rid="ref32">Ozimek and Hanaka, 2021</xref>; <xref ref-type="bibr" rid="ref47">Wu et al., 2022</xref>). More importantly, functional predictions indicated that SBFSF enhanced multiple metabolic pathways including electron transfer, respiration, vitamin biosynthesis, and secondary metabolite production. These improvements indicate a more metabolically active fungal community, which is capable to sustain soil ecosystem functioning.</p>
<p>The differing responses of microbial community to mulching practices indicates a balance between diversity and function maintenance. Even if SM and SBF promoted taxonomic diversity, SBFSF generated a functionally specialized community with greater metabolic potential and lower potential pathogen load. This is indicative that the functional attributes may be more important than overall diversity in supporting soil health and plant productivity.</p>
</sec>
<sec id="sec25">
<label>4.3</label>
<title>Relationships between soil properties, microbial communities and tomato plant growth</title>
<p>Mulching is widely utilized in field crop and vegetable production and has been shown to promote crop growth (<xref ref-type="bibr" rid="ref37">Sharma et al., 2023</xref>; <xref ref-type="bibr" rid="ref40">Wang J. et al., 2025</xref>; <xref ref-type="bibr" rid="ref54">Zhao Y. et al., 2023</xref>). Similarly, our study demonstrated that all mulching treatments significantly enhanced tomato yield compared to CK, with SBFSF exhibiting the most pronounced effect. These improvements can be attributed to the integrated modifications in plant growth and soil microenvironment induced by the different mulching practices.</p>
<p>The higher yield under SBFSF compared to SBF and SM was also consistent with enhanced dry matter accumulation and root growth. Increased root length and area under SBFSF probably contributed to enhanced nutrient and water uptake (<xref ref-type="bibr" rid="ref31">Mooney et al., 2010</xref>; <xref ref-type="bibr" rid="ref16">Guo et al., 2024</xref>) which resulted in higher biomass production and fruit yield. The early advantage in dry matter under SBF at 20&#x202F;days may be attributed to its more effective soil warming during initial growth. However, SBFSF surpassed all other treatments by 40&#x202F;days, indicating that its combined regulation of temperature and moisture better sustained plant development. Furthermore, SBFSF promoted the proliferation of beneficial soil microorganisms such as <italic>Firmicutes</italic>, <italic>Bacillus</italic>, <italic>Lysobacte</italic>, <italic>Ascomycota</italic>, <italic>Mortierellomycota</italic> and <italic>Trichocladium</italic>, consequently, the beneficial microorganisms improved key soil properties, including soil organic matter content, total nitrogen, and available nutrients, ultimately leading to an increasing in tomato production (<xref ref-type="bibr" rid="ref38">Tao et al., 2024</xref>). These findings align with previous reports demonstrating that microbial community modulation can enhance crop productivity (<xref ref-type="bibr" rid="ref42">Wang et al., 2024</xref>; <xref ref-type="bibr" rid="ref22">Li et al., 2020</xref>). Conversely, the stimulation of potential pathogenic species (<italic>Fusarium</italic>, <italic>Alternaria</italic>, <italic>Cladosporium</italic>) under SM may have partially counteracted the benefits of nutrient availability, potentially explaining why tomato production under SM underperformed compared to SBFSF although they had improved some soil properties. Similarly, the SBF treatment also elevated the abundance of potentially pathogenic fungi, including <italic>Cephaliophora</italic> and <italic>Fusarium</italic>. Given its suboptimal soil physicochemical performance, this likely contributed to the reduced yield of SBF compared to SBFSF. Based on partial least squares path modeling (PLS-PM), it suggested that soil properties were strongly and directly associated with tomato growth, whereas mulching practices were linked to enhanced tomato growth primarily through their association with changes in bacterial and fungal diversity and community structure, which in turn were related to soil properties.</p>
<p>Our study revealed an increase in the population of soil potential pathogenic fungi following 3&#x202F;years of plastic film mulching alone and straw mulching alone. Despite this increase, tomato yields remained higher compared to the control. Thus, the potential continuous decrease in tomato yield over successive planting years warrants further observation over an extended period. Our study spanned a limited duration of 3&#x202F;years; moving forward, our research will persist in examining both tomato soil quality and yield. Additionally, we examined soil nutrient and microbial conditions only before tomato vine removal and did not track their dynamics across multiple time points. Investigating those temporal changes is a direction for our future research.</p>
</sec>
</sec>
<sec sec-type="conclusions" id="sec26">
<label>5</label>
<title>Conclusion</title>
<p>This three-year study demonstrates that film-straw dual mulching (SBFSF) is the most effective practice among the treatments tested, enhancing tomato yield by optimizing soil moisture, temperature, and nutrient availability. Its superior performance is primarily attributed to the selective enrichment of beneficial functional microorganisms, such as plant growth-promoting rhizobacteria and key taxa involved in carbon-nitrogen cycling, despite reducing overall microbial diversity. In contrast, single mulching practices promoted certain potential pathogenic fungi. The PLS-PM model indicated that mulching directly shape the microbial community, which promotes tomato growth by indirectly altering soil properties. Although the specific contributions of film versus straw warrant further investigation (e.g., using fully covered single-material controls), the SBFSF system as a whole is advocated as a sustainable agricultural strategy for enhancing tomato productivity via the promotion of a beneficial soil micro-ecosystem.</p>
</sec>
</body>
<back>
<sec sec-type="data-availability" id="sec27">
<title>Data availability statement</title>
<p>The datasets presented in this study can be found in online repositories. The names of the repository/repositories and accession number(s) can be found here: <ext-link xlink:href="https://ngdc.cncb.ac.cn/gsa/browse/CRA037497" ext-link-type="uri">https://ngdc.cncb.ac.cn/gsa/browse/CRA037497</ext-link>.</p>
</sec>
<sec sec-type="author-contributions" id="sec28">
<title>Author contributions</title>
<p>XL: Writing &#x2013; review &#x0026; editing, Formal analysis, Supervision, Writing &#x2013; original draft, Conceptualization, Methodology, Visualization, Investigation. JC: Writing &#x2013; original draft, Investigation, Formal analysis, Data curation. DL: Methodology, Writing &#x2013; original draft, Visualization. KJ: Investigation, Writing &#x2013; original draft. YL: Methodology, Writing &#x2013; original draft. WL: Formal analysis, Writing &#x2013; review &#x0026; editing, Investigation, Conceptualization.</p>
</sec>
<sec sec-type="COI-statement" id="sec29">
<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 sec-type="ai-statement" id="sec30">
<title>Generative AI statement</title>
<p>The author(s) declared that Generative AI was not used in the creation of this manuscript.</p>
<p>Any alternative text (alt text) provided alongside figures in this article has been generated by Frontiers with the support of artificial intelligence and reasonable efforts have been made to ensure accuracy, including review by the authors wherever possible. If you identify any issues, please contact us.</p>
</sec>
<sec sec-type="disclaimer" id="sec31">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<sec sec-type="supplementary-material" id="sec32">
<title>Supplementary material</title>
<p>The Supplementary material for this article can be found online at: <ext-link xlink:href="https://www.frontiersin.org/articles/10.3389/fmicb.2025.1734062/full#supplementary-material" ext-link-type="uri">https://www.frontiersin.org/articles/10.3389/fmicb.2025.1734062/full#supplementary-material</ext-link></p>
<supplementary-material xlink:href="Table_1.docx" id="SM1" mimetype="application/vnd.openxmlformats-officedocument.wordprocessingml.document" xmlns:xlink="http://www.w3.org/1999/xlink"/>
<supplementary-material xlink:href="Data_Sheet_1.docx" id="SM2" mimetype="application/vnd.openxmlformats-officedocument.wordprocessingml.document" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</sec>
<ref-list>
<title>References</title>
<ref id="ref1"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Afoshin</surname><given-names>A. S.</given-names></name> <name><surname>Konstantinov</surname><given-names>M. A.</given-names></name> <name><surname>Toropygin</surname><given-names>I. Y.</given-names></name> <name><surname>Kudryakova</surname><given-names>I. V.</given-names></name> <name><surname>Vasilyeva</surname><given-names>N. V.</given-names></name></person-group> (<year>2020</year>). <article-title>&#x0392;-lytic protease of <italic>Lysobacter capsici</italic> VKM B-2533T</article-title>. <source>Antibiotics</source> <volume>9</volume>:<fpage>744</fpage>. doi: <pub-id pub-id-type="doi">10.3390/antibiotics9110744</pub-id>, <pub-id pub-id-type="pmid">33126447</pub-id></mixed-citation></ref>
<ref id="ref2"><mixed-citation publication-type="book"><person-group person-group-type="author"><name><surname>Bao</surname><given-names>S. D.</given-names></name></person-group> (<year>2000</year>). <source>Soil and agricultural chemical analysis</source>. <edition>3rd</edition> Edn. <publisher-loc>Beijing</publisher-loc>: <publisher-name>China Agricultural Press</publisher-name>.</mixed-citation></ref>
<ref id="ref3"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Biswal</surname><given-names>P.</given-names></name> <name><surname>Swain</surname><given-names>D. K.</given-names></name> <name><surname>Jha</surname><given-names>M. K.</given-names></name></person-group> (<year>2022</year>). <article-title>Straw mulch with limited drip irrigation influenced soil microclimate in improving tuber yield and water productivity of potato in subtropical India</article-title>. <source>Soil Tillage Res.</source> <volume>223</volume>:<fpage>105484</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.still.2022.105484</pub-id></mixed-citation></ref>
<ref id="ref4"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Chai</surname><given-names>Y.</given-names></name> <name><surname>Chai</surname><given-names>Q.</given-names></name> <name><surname>Li</surname><given-names>R.</given-names></name> <name><surname>Li</surname><given-names>Y.</given-names></name> <name><surname>Yang</surname><given-names>C.</given-names></name> <name><surname>Cheng</surname><given-names>H.</given-names></name> <etal/></person-group>. (<year>2022</year>). <article-title>Straw strip mulching in a semiarid rainfed agroecosystem achieves winter wheat yields similar to those of full plastic mulching by optimizing the soil hydrothermal regime</article-title>. <source>Crop J.</source> <volume>10</volume>, <fpage>879</fpage>&#x2013;<lpage>892</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.cj.2021.09.004</pub-id></mixed-citation></ref>
<ref id="ref5"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Chakraborty</surname><given-names>D.</given-names></name> <name><surname>Nagarajan</surname><given-names>S.</given-names></name> <name><surname>Aggarwal</surname><given-names>P.</given-names></name> <name><surname>Gupta</surname><given-names>V. K.</given-names></name> <name><surname>Tomar</surname><given-names>R. K.</given-names></name> <name><surname>Garg</surname><given-names>R. N.</given-names></name> <etal/></person-group>. (<year>2008</year>). <article-title>Effect of mulching on soil and plant water status, and the growth and yield of wheat (<italic>Triticum aestivum L.</italic>) in a semi-arid environment</article-title>. <source>Agric. Water Manag.</source> <volume>95</volume>, <fpage>1323</fpage>&#x2013;<lpage>1334</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.agwat.2008.06.001</pub-id></mixed-citation></ref>
<ref id="ref6"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Chen</surname><given-names>Y. L.</given-names></name> <name><surname>Xin</surname><given-names>L.</given-names></name> <name><surname>Liu</surname><given-names>J. T.</given-names></name> <name><surname>Yuan</surname><given-names>M. Z.</given-names></name> <name><surname>Liu</surname><given-names>S. T.</given-names></name> <name><surname>Jiang</surname><given-names>W.</given-names></name> <etal/></person-group>. (<year>2017</year>). <article-title>Changes in bacterial community of soil induced by long-term straw returning</article-title>. <source>Sci. Agric.</source> <volume>74</volume>, <fpage>349</fpage>&#x2013;<lpage>356</lpage>. doi: <pub-id pub-id-type="doi">10.1590/1678-992X-2016-0025</pub-id></mixed-citation></ref>
<ref id="ref7"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Chen</surname><given-names>W.</given-names></name> <name><surname>Yuan</surname><given-names>W.</given-names></name> <name><surname>Wang</surname><given-names>J.</given-names></name> <name><surname>Wang</surname><given-names>Z.</given-names></name> <name><surname>Zhou</surname><given-names>Z.</given-names></name> <name><surname>Liu</surname><given-names>S.</given-names></name></person-group> (<year>2022</year>). <article-title>No-tillage combined with appropriate amount of straw returning increased soil biochemical properties</article-title>. <source>Sustainability</source> <volume>14</volume>:<fpage>4875</fpage>. doi: <pub-id pub-id-type="doi">10.3390/su14094875</pub-id></mixed-citation></ref>
<ref id="ref8"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Dai</surname><given-names>Z.</given-names></name> <name><surname>Fan</surname><given-names>J.</given-names></name> <name><surname>Fu</surname><given-names>W.</given-names></name> <name><surname>Niu</surname><given-names>X.</given-names></name> <name><surname>Yang</surname><given-names>Q.</given-names></name> <name><surname>Hao</surname><given-names>M.</given-names></name></person-group> (<year>2024</year>). <article-title>Changes in soil microbial community and co-occurrence network after long-term no-tillage and mulching in dryland farming</article-title>. <source>Plant Soil</source> <volume>495</volume>, <fpage>201</fpage>&#x2013;<lpage>220</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s11104-023-06315-w</pub-id></mixed-citation></ref>
<ref id="ref9"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Dang</surname><given-names>K.</given-names></name> <name><surname>Liang</surname><given-names>H.</given-names></name> <name><surname>Guo</surname><given-names>S.</given-names></name> <name><surname>Fan</surname><given-names>Z.</given-names></name> <name><surname>Li</surname><given-names>H.</given-names></name> <name><surname>Ma</surname><given-names>M.</given-names></name> <etal/></person-group>. (<year>2025</year>). <article-title>Adaptive regulation of microbial community characteristics in response to nutrient limitations under mulching practices across distinct climate zones</article-title>. <source>Soil Tillage Res.</source> <volume>248</volume>:<fpage>106465</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.still.2025.106465</pub-id></mixed-citation></ref>
<ref id="ref10"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Dong</surname><given-names>W.</given-names></name> <name><surname>Si</surname><given-names>P.</given-names></name> <name><surname>Liu</surname><given-names>E.</given-names></name> <name><surname>Yan</surname><given-names>C.</given-names></name> <name><surname>Zhang</surname><given-names>Z.</given-names></name> <name><surname>Zhang</surname><given-names>Y.</given-names></name></person-group> (<year>2017</year>). <article-title>Influence of film mulching on soil microbial community in a rainfed region of northeastern China</article-title>. <source>Sci. Rep.</source> <volume>7</volume>:<fpage>8468</fpage>. doi: <pub-id pub-id-type="doi">10.1038/s41598-017-08575-w</pub-id>, <pub-id pub-id-type="pmid">28814759</pub-id></mixed-citation></ref>
<ref id="ref11"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Du</surname><given-names>X.</given-names></name></person-group> (<year>2024</year>). <article-title>Improving saline-alkali soil with agricultural waste in China: a review</article-title>. <source>Commun. Soil Sci. Plant Anal.</source> <volume>55</volume>, <fpage>2651</fpage>&#x2013;<lpage>2665</lpage>. doi: <pub-id pub-id-type="doi">10.1080/00103624.2024.2372402</pub-id></mixed-citation></ref>
<ref id="ref13"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Fu</surname><given-names>X.</given-names></name> <name><surname>Wang</surname><given-names>J.</given-names></name> <name><surname>Sainju</surname><given-names>U. M.</given-names></name> <name><surname>Zhao</surname><given-names>F.</given-names></name> <name><surname>Liu</surname><given-names>W.</given-names></name></person-group> (<year>2019</year>). <article-title>Soil microbial community and carbon and nitrogen fractions responses to mulching under winter wheat</article-title>. <source>Appl. Soil Ecol.</source> <volume>139</volume>, <fpage>64</fpage>&#x2013;<lpage>68</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.apsoil.2019.03.018</pub-id></mixed-citation></ref>
<ref id="ref14"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Gon&#x00E7;alves</surname><given-names>O. S.</given-names></name> <name><surname>Fernandes</surname><given-names>A. S.</given-names></name> <name><surname>Tupy</surname><given-names>S. M.</given-names></name> <name><surname>Ferreira</surname><given-names>T. G.</given-names></name> <name><surname>Almeida</surname><given-names>L. N.</given-names></name> <name><surname>Creevey</surname><given-names>C. J.</given-names></name> <etal/></person-group>. (<year>2024</year>). <article-title>Insights into plant interactions and the biogeochemical role of the globally widespread Acidobacteriota phylum</article-title>. <source>Soil Biol. Biochem.</source> <volume>192</volume>:<fpage>109369</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.soilbio.2024.109369</pub-id></mixed-citation></ref>
<ref id="ref15"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Gu</surname><given-names>X.</given-names></name> <name><surname>Cai</surname><given-names>H.</given-names></name> <name><surname>Fang</surname><given-names>H.</given-names></name> <name><surname>Li</surname><given-names>Y.</given-names></name> <name><surname>Chen</surname><given-names>P.</given-names></name> <name><surname>Li</surname><given-names>Y.</given-names></name></person-group> (<year>2020</year>). <article-title>Effects of degradable film mulching on crop yield and water use efficiency in China: a meta-analysis</article-title>. <source>Soil Tillage Res.</source> <volume>202</volume>:<fpage>104676</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.still.2020.104676</pub-id></mixed-citation></ref>
<ref id="ref16"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Guo</surname><given-names>C.</given-names></name> <name><surname>Bao</surname><given-names>X.</given-names></name> <name><surname>Sun</surname><given-names>H.</given-names></name> <name><surname>Zhu</surname><given-names>L.</given-names></name> <name><surname>Zhang</surname><given-names>Y.</given-names></name> <name><surname>Zhang</surname><given-names>K.</given-names></name> <etal/></person-group>. (<year>2024</year>). <article-title>Optimizing root system architecture to improve cotton drought tolerance and minimize yield loss during mild drought stress</article-title>. <source>Field Crop Res.</source> <volume>308</volume>:<fpage>109305</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.fcr.2024.109305</pub-id></mixed-citation></ref>
<ref id="ref17"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Han</surname><given-names>Y.</given-names></name> <name><surname>Lu</surname><given-names>L.</given-names></name> <name><surname>Wang</surname><given-names>L.</given-names></name> <name><surname>Liu</surname><given-names>Z. T.</given-names></name> <name><surname>Huang</surname><given-names>P. F.</given-names></name> <name><surname>Chen</surname><given-names>S. C.</given-names></name> <etal/></person-group>. (<year>2024</year>). <article-title>In-situ straw return, combined with plastic film use, influences soil properties and tomato quality and yield in greenhouse conditions</article-title>. <source>Agric. Commun.</source> <volume>2</volume>:<fpage>100028</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.agrcom.2024.100028</pub-id></mixed-citation></ref>
<ref id="ref18"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Hu</surname><given-names>C.</given-names></name> <name><surname>Lei</surname><given-names>F.</given-names></name> <name><surname>Zhang</surname><given-names>X.</given-names></name> <name><surname>Shi</surname><given-names>J.</given-names></name> <name><surname>Li</surname><given-names>J.</given-names></name> <name><surname>Zuo</surname><given-names>X.</given-names></name> <etal/></person-group>. (<year>2023</year>). <article-title>Black carbon derived from pyrolysis of maize straw and polystyrene microplastics affects soil biodiversity</article-title>. <source>Sci. Total Environ.</source> <volume>881</volume>:<fpage>163398</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.scitotenv.2023.163398</pub-id>, <pub-id pub-id-type="pmid">37061062</pub-id></mixed-citation></ref>
<ref id="ref19"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Hu</surname><given-names>W.</given-names></name> <name><surname>Li</surname><given-names>Q.</given-names></name> <name><surname>Wang</surname><given-names>W.</given-names></name> <name><surname>Lin</surname><given-names>X.</given-names></name> <name><surname>He</surname><given-names>Z.</given-names></name> <name><surname>Li</surname><given-names>G.</given-names></name></person-group> (<year>2024</year>). <article-title>Straw mulching decreased the contribution of Fe-bound organic carbon to soil organic carbon in a banana orchard</article-title>. <source>Appl. Soil Ecol.</source> <volume>194</volume>:<fpage>105177</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.apsoil.2023.105177</pub-id></mixed-citation></ref>
<ref id="ref20"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Kong</surname><given-names>H.</given-names></name> <name><surname>Fu</surname><given-names>X. C.</given-names></name> <name><surname>Chen</surname><given-names>X.</given-names></name> <name><surname>Ding</surname><given-names>Z. M.</given-names></name> <name><surname>Yang</surname><given-names>Y.</given-names></name> <name><surname>Xu</surname><given-names>H. W.</given-names></name> <etal/></person-group>. (<year>2023</year>). <article-title>The ester derivatives of ferulic acid exhibit strong inhibitory effect on the growth of <italic>Alternaria alternata</italic> in vitro and in vivo</article-title>. <source>Postharvest Biol. Technol.</source> <volume>196</volume>:<fpage>112158</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.postharvbio.2022.112158</pub-id></mixed-citation></ref>
<ref id="ref21"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Li</surname><given-names>D. Y.</given-names></name> <name><surname>Yu</surname><given-names>Z.</given-names></name> <name><surname>Li</surname><given-names>Y. Y.</given-names></name> <name><surname>Li</surname><given-names>X. Q.</given-names></name> <name><surname>Li</surname><given-names>Y. Q.</given-names></name> <name><surname>Li</surname><given-names>B. L.</given-names></name> <etal/></person-group>. (<year>2024</year>). <article-title>Effects of combined microplastic and cadmium pollution on sorghum growth, cd accumulation, and rhizosphere microbial functions</article-title>. <source>Ecotoxicol. Environ. Saf.</source> <volume>277</volume>:<fpage>116380</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.ecoenv.2024.116380</pub-id></mixed-citation></ref>
<ref id="ref22"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Li</surname><given-names>F.</given-names></name> <name><surname>Zhang</surname><given-names>S.</given-names></name> <name><surname>Wang</surname><given-names>Y.</given-names></name> <name><surname>Li</surname><given-names>Y.</given-names></name> <name><surname>Li</surname><given-names>P.</given-names></name> <name><surname>Chen</surname><given-names>L.</given-names></name> <etal/></person-group>. (<year>2020</year>). <article-title>Rare fungus, <italic>Mortierella capitata</italic>, promotes crop growth by stimulating primary metabolisms related genes and reshaping rhizosphere bacterial community</article-title>. <source>Soil Biol. Biochem.</source> <volume>151</volume>:<fpage>108017</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.soilbio.2020.108017</pub-id></mixed-citation></ref>
<ref id="ref23"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Ling</surname><given-names>N.</given-names></name> <name><surname>Zhu</surname><given-names>C.</given-names></name> <name><surname>Xue</surname><given-names>C.</given-names></name> <name><surname>Chen</surname><given-names>H.</given-names></name> <name><surname>Duan</surname><given-names>Y.</given-names></name> <name><surname>Peng</surname><given-names>C.</given-names></name> <etal/></person-group>. (<year>2016</year>). <article-title>Insight into how organic amendments can shape the soil microbiome in long-term field experiments as revealed by network analysis</article-title>. <source>Soil Biol. Biochem.</source> <volume>99</volume>, <fpage>137</fpage>&#x2013;<lpage>149</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.soilbio.2016.05.005</pub-id></mixed-citation></ref>
<ref id="ref24"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Liu</surname><given-names>Q.</given-names></name> <name><surname>Tang</surname><given-names>L.</given-names></name> <name><surname>Sun</surname><given-names>H.</given-names></name> <name><surname>Kong</surname><given-names>X.</given-names></name> <name><surname>Jiao</surname><given-names>Y.</given-names></name> <name><surname>Wu</surname><given-names>W.</given-names></name> <etal/></person-group>. (<year>2024</year>). <article-title>Responses of the fungal-bacterial community and network to surface mulching and nitrogen fertilization in the loess plateau</article-title>. <source>Plant Soil</source> <volume>494</volume>, <fpage>111</fpage>&#x2013;<lpage>126</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s11104-023-06260-8</pub-id></mixed-citation></ref>
<ref id="ref25"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Liu</surname><given-names>E.</given-names></name> <name><surname>Teclemariam</surname><given-names>S. G.</given-names></name> <name><surname>Yan</surname><given-names>C.</given-names></name> <name><surname>Yu</surname><given-names>J.</given-names></name> <name><surname>Gu</surname><given-names>R.</given-names></name> <name><surname>Liu</surname><given-names>S.</given-names></name> <etal/></person-group>. (<year>2014</year>). <article-title>Long-term effects of no-tillage management practice on soil organic carbon and its fractions in the northern China</article-title>. <source>Geoderma</source> <volume>213</volume>, <fpage>379</fpage>&#x2013;<lpage>384</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.geoderma.2013.08.021</pub-id></mixed-citation></ref>
<ref id="ref26"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Luo</surname><given-names>D.</given-names></name> <name><surname>Huang</surname><given-names>H. Z.</given-names></name> <name><surname>Wang</surname><given-names>J. H.</given-names></name> <name><surname>Wang</surname><given-names>T. F.</given-names></name> <name><surname>Zhang</surname><given-names>Q. F.</given-names></name> <name><surname>Wang</surname><given-names>Y. G.</given-names></name> <etal/></person-group>. (<year>2025</year>). <article-title>Inoculations of phosphorus-solubilizing bacteria alter soil properties, microbial communities, and improve crop yield and fruit quality in the tomato production system</article-title>. <source>Appl. Soil Ecol.</source> <volume>207</volume>:<fpage>105944</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.apsoil.2025.105944</pub-id></mixed-citation></ref>
<ref id="ref27"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Luo</surname><given-names>Z.</given-names></name> <name><surname>Ma</surname><given-names>J.</given-names></name> <name><surname>Chen</surname><given-names>F.</given-names></name> <name><surname>Li</surname><given-names>X.</given-names></name> <name><surname>Hou</surname><given-names>H.</given-names></name> <name><surname>Zhang</surname><given-names>S.</given-names></name></person-group> (<year>2019</year>). <article-title>Cracks reinforce the interactions among soil bacterial communities in the coal mining area of loess plateau, China</article-title>. <source>Int. J. Environ. Res. Public Health</source> <volume>16</volume>:<fpage>4892</fpage>. doi: <pub-id pub-id-type="doi">10.3390/ijerph16244892</pub-id>, <pub-id pub-id-type="pmid">31817220</pub-id></mixed-citation></ref>
<ref id="ref28"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Luo</surname><given-names>C. L.</given-names></name> <name><surname>Zhang</surname><given-names>X. F.</given-names></name> <name><surname>Duan</surname><given-names>H. X.</given-names></name> <name><surname>Mburu</surname><given-names>D. M.</given-names></name> <name><surname>Ren</surname><given-names>H. X.</given-names></name> <name><surname>Kavagi</surname><given-names>L.</given-names></name> <etal/></person-group>. (<year>2020</year>). <article-title>Film-straw dual mulching boosts wheat productivity and soil quality under the El Ni&#x00F1;o in semiarid Kenya</article-title>. <source>Sci. Total Environ.</source> <volume>738</volume>:<fpage>139808</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.scitotenv.2020.139808</pub-id></mixed-citation></ref>
<ref id="ref29"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Ma</surname><given-names>S.</given-names></name> <name><surname>Fang</surname><given-names>C.</given-names></name> <name><surname>Sun</surname><given-names>X.</given-names></name> <name><surname>Han</surname><given-names>L.</given-names></name> <name><surname>He</surname><given-names>X.</given-names></name> <name><surname>Huang</surname><given-names>G.</given-names></name></person-group> (<year>2018</year>). <article-title>Bacterial community succession during pig manure and wheat straw aerobic composting covered with a semi-permeable membrane under slight positive pressure</article-title>. <source>Bioresour. Technol.</source> <volume>259</volume>, <fpage>221</fpage>&#x2013;<lpage>227</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.biortech.2018.03.054</pub-id>, <pub-id pub-id-type="pmid">29558720</pub-id></mixed-citation></ref>
<ref id="ref30"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Miao</surname><given-names>F.</given-names></name> <name><surname>Pu</surname><given-names>X.</given-names></name> <name><surname>Hou</surname><given-names>X.</given-names></name> <name><surname>Li</surname><given-names>R.</given-names></name></person-group> (<year>2025</year>). <article-title>Enhancing soil hydrothermal conditions and improving potato yields through furrow&#x2013;ridge mulching under drought&#x2013;prone semi&#x2013;humid climatic conditions</article-title>. <source>Agric. Water Manag.</source> <volume>319</volume>:<fpage>109769</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.agwat.2025.109769</pub-id></mixed-citation></ref>
<ref id="ref31"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Mooney</surname><given-names>K. A.</given-names></name> <name><surname>Halitschke</surname><given-names>R.</given-names></name> <name><surname>Kessler</surname><given-names>A.</given-names></name> <name><surname>Agrawal</surname><given-names>A. A.</given-names></name></person-group> (<year>2010</year>). <article-title>Evolutionary trade-offs in plants mediate the strength of trophic cascades</article-title>. <source>Science</source> <volume>327</volume>, <fpage>1642</fpage>&#x2013;<lpage>1644</lpage>. doi: <pub-id pub-id-type="doi">10.1126/science.1184814</pub-id></mixed-citation></ref>
<ref id="ref32"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Ozimek</surname><given-names>E.</given-names></name> <name><surname>Hanaka</surname><given-names>A.</given-names></name></person-group> (<year>2021</year>). <article-title>Mortierella species as the plant growth-promoting fungi present in the agricultural soils</article-title>. <source>Agriculture</source> <volume>11</volume>:<fpage>7</fpage>. doi: <pub-id pub-id-type="doi">10.3390/agriculture11010007</pub-id></mixed-citation></ref>
<ref id="ref33"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Philippot</surname><given-names>L.</given-names></name> <name><surname>Chenu</surname><given-names>C.</given-names></name> <name><surname>Kappler</surname><given-names>A.</given-names></name> <name><surname>Rillig</surname><given-names>M. C.</given-names></name> <name><surname>Fierer</surname><given-names>N.</given-names></name></person-group> (<year>2023</year>). <article-title>The interplay between microbial communities and soil properties</article-title>. <source>Nat. Rev. Microbiol.</source> <volume>22</volume>, <fpage>226</fpage>&#x2013;<lpage>239</lpage>. doi: <pub-id pub-id-type="doi">10.1038/s41579-023-00980-5</pub-id>, <pub-id pub-id-type="pmid">37863969</pub-id></mixed-citation></ref>
<ref id="ref34"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Qi</surname><given-names>Y.</given-names></name> <name><surname>Ossowicki</surname><given-names>A.</given-names></name> <name><surname>Yergeau</surname><given-names>&#x00C9;.</given-names></name> <name><surname>Vigani</surname><given-names>G.</given-names></name> <name><surname>Geissen</surname><given-names>V.</given-names></name> <name><surname>Garbeva</surname><given-names>P.</given-names></name></person-group> (<year>2022</year>). <article-title>Plastic mulch film residues in agriculture: impact on soil suppressiveness, plant growth, and microbial communities</article-title>. <source>FEMS Microbiol. Ecol.</source> <volume>98</volume>:<fpage>fiac017</fpage>. doi: <pub-id pub-id-type="doi">10.1093/femsec/fiac017</pub-id>, <pub-id pub-id-type="pmid">35150249</pub-id></mixed-citation></ref>
<ref id="ref35"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Qu</surname><given-names>Y.</given-names></name> <name><surname>Feng</surname><given-names>B.</given-names></name></person-group> (<year>2022</year>). <article-title>Straw mulching improved yield of field buckwheat (<italic>Fagopyrum</italic>) by increasing water-temperature use and soil carbon in rain-fed farmland</article-title>. <source>Acta Ecol. Sin.</source> <volume>42</volume>, <fpage>11</fpage>&#x2013;<lpage>16</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.chnaes.2020.11.008</pub-id></mixed-citation></ref>
<ref id="ref36"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Saxena</surname><given-names>A. K.</given-names></name> <name><surname>Kumar</surname><given-names>M.</given-names></name> <name><surname>Chakdar</surname><given-names>H.</given-names></name> <name><surname>Anuroopa</surname><given-names>N.</given-names></name> <name><surname>Bagyaraj</surname><given-names>D. J.</given-names></name></person-group> (<year>2020</year>). <article-title>Bacillus species in soil as a natural resource for plant health and nutrition</article-title>. <source>J. Appl. Microbiol.</source> <volume>128</volume>, <fpage>1583</fpage>&#x2013;<lpage>1594</lpage>. doi: <pub-id pub-id-type="doi">10.1111/jam.14506</pub-id>, <pub-id pub-id-type="pmid">31705597</pub-id></mixed-citation></ref>
<ref id="ref37"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Sharma</surname><given-names>S.</given-names></name> <name><surname>Basnet</surname><given-names>B.</given-names></name> <name><surname>Bhattarai</surname><given-names>K.</given-names></name> <name><surname>Sedhai</surname><given-names>A.</given-names></name> <name><surname>Khanal</surname><given-names>K.</given-names></name></person-group> (<year>2023</year>). <article-title>The influence of different mulching materials on tomato&#x2019;s vegetative, reproductive, and yield in Dhankuta, Nepal</article-title>. <source>J. Agric. Food Res.</source> <volume>11</volume>:<fpage>100463</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.jafr.2022.100463</pub-id></mixed-citation></ref>
<ref id="ref38"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Tao</surname><given-names>W. Q.</given-names></name> <name><surname>Wu</surname><given-names>Q. Q.</given-names></name> <name><surname>Zhang</surname><given-names>J.</given-names></name> <name><surname>Chang</surname><given-names>T. T.</given-names></name> <name><surname>Liu</surname><given-names>X. N.</given-names></name></person-group> (<year>2024</year>). <article-title>Effects of applying organic amendments on soil aggregate structure and tomato yield in facility agriculture</article-title>. <source>Plants</source> <volume>13</volume>:<fpage>3064</fpage>. doi: <pub-id pub-id-type="doi">10.3390/plants13213064</pub-id>, <pub-id pub-id-type="pmid">39519978</pub-id></mixed-citation></ref>
<ref id="ref39"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Wang</surname><given-names>J.</given-names></name> <name><surname>Chapman</surname><given-names>S. J.</given-names></name> <name><surname>Yao</surname><given-names>H.</given-names></name></person-group> (<year>2015</year>). <article-title>The effect of storage on microbial activity and bacterial community structure of drained and flooded paddy soil</article-title>. <source>J. Soils Sediments</source> <volume>15</volume>, <fpage>880</fpage>&#x2013;<lpage>889</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s11368-014-1053-7</pub-id></mixed-citation></ref>
<ref id="ref40"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Wang</surname><given-names>J.</given-names></name> <name><surname>Fan</surname><given-names>J.</given-names></name> <name><surname>Wang</surname><given-names>H.</given-names></name> <name><surname>Wang</surname><given-names>X.</given-names></name> <name><surname>Xing</surname><given-names>Y.</given-names></name> <name><surname>Gao</surname><given-names>Y.</given-names></name> <etal/></person-group>. (<year>2025</year>). <article-title>Dual-mulching under no-tillage promotes maize root growth and improves yield by optimizing soil hydrothermal conditions in semi-arid regions</article-title>. <source>Agric. Water Manag.</source> <volume>312</volume>:<fpage>109428</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.agwat.2025.109428</pub-id></mixed-citation></ref>
<ref id="ref41"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Wang</surname><given-names>Y.</given-names></name> <name><surname>Pang</surname><given-names>J.</given-names></name> <name><surname>Zhang</surname><given-names>M.</given-names></name> <name><surname>Tian</surname><given-names>Z.</given-names></name> <name><surname>Wei</surname><given-names>T.</given-names></name> <name><surname>Jia</surname><given-names>Z.</given-names></name> <etal/></person-group>. (<year>2023</year>). <article-title>Is adding biochar be better than crop straw for improving soil aggregates stability and organic carbon contents in film mulched fields in semiarid regions?&#x2014;evidence of 5-year field experiment</article-title>. <source>J. Environ. Manag.</source> <volume>338</volume>:<fpage>117711</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.jenvman.2023.117711</pub-id>, <pub-id pub-id-type="pmid">36996557</pub-id></mixed-citation></ref>
<ref id="ref42"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Wang</surname><given-names>X.</given-names></name> <name><surname>Qian</surname><given-names>R.</given-names></name> <name><surname>Han</surname><given-names>Y.</given-names></name> <name><surname>Ji</surname><given-names>Z.</given-names></name> <name><surname>Yang</surname><given-names>Q.</given-names></name> <name><surname>Wang</surname><given-names>L.</given-names></name> <etal/></person-group>. (<year>2024</year>). <article-title>Straw return can increase maize yield by regulating soil bacteria and improving soil properties in arid and semi-arid areas</article-title>. <source>Eur. J. Agron.</source> <volume>161</volume>:<fpage>127389</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.eja.2024.127389</pub-id></mixed-citation></ref>
<ref id="ref43"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Wang</surname><given-names>J.</given-names></name> <name><surname>Shi</surname><given-names>X.</given-names></name> <name><surname>Li</surname><given-names>Z.</given-names></name> <name><surname>Zhang</surname><given-names>Y.</given-names></name> <name><surname>Liu</surname><given-names>Y.</given-names></name> <name><surname>Peng</surname><given-names>Y.</given-names></name></person-group> (<year>2021</year>). <article-title>Responses of runoff and soil erosion to planting pattern, row direction, and straw mulching on sloped farmland in the corn belt of Northeast China</article-title>. <source>Agric. Water Manag.</source> <volume>253</volume>:<fpage>106935</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.agwat.2021.106935</pub-id></mixed-citation></ref>
<ref id="ref44"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Wang</surname><given-names>F.</given-names></name> <name><surname>Xie</surname><given-names>R.</given-names></name> <name><surname>Ming</surname><given-names>B.</given-names></name> <name><surname>Wang</surname><given-names>K.</given-names></name> <name><surname>Hou</surname><given-names>P.</given-names></name> <name><surname>Chen</surname><given-names>J.</given-names></name> <etal/></person-group>. (<year>2021</year>). <article-title>Dry matter accumulation after silking and kernel weight are the key factors for increasing maize yield and water use efficiency</article-title>. <source>Agric. Water Manag.</source> <volume>254</volume>:<fpage>106938</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.agwat.2021.106938</pub-id></mixed-citation></ref>
<ref id="ref45"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Wang</surname><given-names>Y.</given-names></name> <name><surname>Yu</surname><given-names>A.</given-names></name> <name><surname>Wang</surname><given-names>P.</given-names></name> <name><surname>Shang</surname><given-names>Y.</given-names></name> <name><surname>Wang</surname><given-names>F.</given-names></name> <name><surname>Lyu</surname><given-names>H.</given-names></name> <etal/></person-group>. (<year>2025</year>). <article-title>No-tillage with total green manure mulching can improve soil moisture and temperature environment, promote maize root structure and photosynthetic capacity to increase maize yield</article-title>. <source>J. Integr. Agric.</source> 24, 4211&#x2013;4224. doi: <pub-id pub-id-type="doi">10.1016/j.jia.2025.03.010</pub-id></mixed-citation></ref>
<ref id="ref46"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Wr&#x00F3;blewska</surname><given-names>K.</given-names></name> <name><surname>Szumny</surname><given-names>A.</given-names></name> <name><surname>&#x017B;arowska</surname><given-names>B.</given-names></name> <name><surname>Kromer</surname><given-names>K.</given-names></name> <name><surname>D&#x0119;bicz</surname><given-names>R.</given-names></name> <name><surname>Fabian</surname><given-names>S.</given-names></name></person-group> (<year>2019</year>). <article-title>Impact of mulching on growth essential oil composition and its biological activity in <italic>Monarda didyma</italic> L</article-title>. <source>Ind. Crop. Prod.</source> <volume>129</volume>, <fpage>299</fpage>&#x2013;<lpage>308</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.indcrop.2018.11.076</pub-id></mixed-citation></ref>
<ref id="ref47"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Wu</surname><given-names>J.</given-names></name> <name><surname>Shi</surname><given-names>Z.</given-names></name> <name><surname>Zhu</surname><given-names>J.</given-names></name> <name><surname>Cao</surname><given-names>A.</given-names></name> <name><surname>Fang</surname><given-names>W.</given-names></name> <name><surname>Yan</surname><given-names>D.</given-names></name> <etal/></person-group>. (<year>2022</year>). <article-title>Taxonomic response of bacterial and fungal populations to biofertilizers applied to soil or substrate in greenhouse-grown cucumber</article-title>. <source>Sci. Rep.</source> <volume>12</volume>:<fpage>18522</fpage>. doi: <pub-id pub-id-type="doi">10.1038/s41598-022-22673-4</pub-id>, <pub-id pub-id-type="pmid">36323754</pub-id></mixed-citation></ref>
<ref id="ref48"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Yang</surname><given-names>F.</given-names></name> <name><surname>He</surname><given-names>B.</given-names></name> <name><surname>Dong</surname><given-names>B.</given-names></name> <name><surname>Zhang</surname><given-names>G.</given-names></name></person-group> (<year>2024</year>). <article-title>Film-straw dual mulching improves soil fertility and maize yield in dryland farming by increasing straw-degrading bacterial abundance and their positive cooperation</article-title>. <source>Agric. Ecosyst. Environ.</source> <volume>367</volume>:<fpage>108997</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.agee.2024.108997</pub-id></mixed-citation></ref>
<ref id="ref49"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Zhang</surname><given-names>M.</given-names></name> <name><surname>Dang</surname><given-names>P.</given-names></name> <name><surname>Haegeman</surname><given-names>B.</given-names></name> <name><surname>Han</surname><given-names>X.</given-names></name> <name><surname>Wang</surname><given-names>X.</given-names></name> <name><surname>Pu</surname><given-names>X.</given-names></name> <etal/></person-group>. (<year>2024</year>). <article-title>The effects of straw return on soil bacterial diversity and functional profiles: a meta-analysis</article-title>. <source>Soil Biol. Biochem.</source> <volume>195</volume>:<fpage>109484</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.soilbio.2024.109484</pub-id></mixed-citation></ref>
<ref id="ref50"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Zhang</surname><given-names>X.</given-names></name> <name><surname>Liu</surname><given-names>H.</given-names></name> <name><surname>Xie</surname><given-names>X.</given-names></name> <name><surname>Sun</surname><given-names>M.</given-names></name> <name><surname>Du</surname><given-names>N.</given-names></name> <name><surname>Xue</surname><given-names>D.</given-names></name></person-group> (<year>2025</year>). <article-title>Physiological and transcriptomic analysis of tomato leaf mold resistance gene Cf-12 against <italic>Cladosporium fulvum</italic></article-title>. <source>Physiol. Mol. Plant Pathol.</source> <volume>138</volume>:<fpage>102692</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.pmpp.2025.102692</pub-id></mixed-citation></ref>
<ref id="ref51"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Zhang</surname><given-names>Y.</given-names></name> <name><surname>Wang</surname><given-names>J.</given-names></name> <name><surname>Gong</surname><given-names>S.</given-names></name> <name><surname>Xu</surname><given-names>D.</given-names></name> <name><surname>Mo</surname><given-names>Y.</given-names></name> <name><surname>Zhang</surname><given-names>B.</given-names></name></person-group> (<year>2021</year>). <article-title>Straw mulching improves soil water content, increases flag leaf photosynthetic parameters and maintains the yield of winter wheat with different irrigation amounts</article-title>. <source>Agric. Water Manag.</source> <volume>249</volume>:<fpage>106809</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.agwat.2021.106809</pub-id></mixed-citation></ref>
<ref id="ref52"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Zhao</surname><given-names>P. P.</given-names></name> <name><surname>Huang</surname><given-names>Y. T.</given-names></name> <name><surname>Liu</surname><given-names>B. Y.</given-names></name> <name><surname>Chen</surname><given-names>J. Y.</given-names></name> <name><surname>Lei</surname><given-names>Z. Y.</given-names></name> <name><surname>Zhang</surname><given-names>Y. H.</given-names></name> <etal/></person-group>. (<year>2024</year>). <article-title>Effects of daytime and nighttime warming on soil microbial diversity</article-title>. <source>Geoderma</source> <volume>447</volume>:<fpage>116909</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.geoderma.2024.116909</pub-id></mixed-citation></ref>
<ref id="ref53"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Zhao</surname><given-names>S. Q.</given-names></name> <name><surname>Li</surname><given-names>J.</given-names></name> <name><surname>Liu</surname><given-names>J. P.</given-names></name> <name><surname>Xiao</surname><given-names>S. Y.</given-names></name> <name><surname>Yang</surname><given-names>S. M.</given-names></name> <name><surname>Mei</surname><given-names>J. H.</given-names></name> <etal/></person-group>. (<year>2023</year>). <article-title>Secondary metabolites of Alternaria: a comprehensive review of chemical diversity and pharmacological properties</article-title>. <source>Front. Microbiol.</source> <volume>13</volume>:<fpage>1085666</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fmicb.2022.1085666</pub-id>, <pub-id pub-id-type="pmid">36687635</pub-id></mixed-citation></ref>
<ref id="ref54"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Zhao</surname><given-names>Y.</given-names></name> <name><surname>Mao</surname><given-names>X.</given-names></name> <name><surname>Li</surname><given-names>S.</given-names></name> <name><surname>Huang</surname><given-names>X.</given-names></name> <name><surname>Che</surname><given-names>J.</given-names></name> <name><surname>Ma</surname><given-names>C.</given-names></name></person-group> (<year>2023</year>). <article-title>A review of plastic film mulching on water, heat, nitrogen balance, and crop growth in farmland in China</article-title>. <source>Agronomy</source> <volume>13</volume>:<fpage>2515</fpage>. doi: <pub-id pub-id-type="doi">10.3390/agronomy13102515</pub-id></mixed-citation></ref>
<ref id="ref55"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Zhao</surname><given-names>L.</given-names></name> <name><surname>Yang</surname><given-names>P.</given-names></name> <name><surname>Zhao</surname><given-names>Z.</given-names></name> <name><surname>Xia</surname><given-names>Y.</given-names></name></person-group> (<year>2023</year>). <article-title>First report of <italic>Cephaliophora tropica</italic> causing leaf spot on tomato (<italic>Solanum lycopersicum</italic> cv. Picus) in USA</article-title>. <source>Crop Prot.</source> <volume>172</volume>:<fpage>106324</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.cropro.2023.106324</pub-id></mixed-citation></ref>
<ref id="ref56"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Zheng</surname><given-names>W.</given-names></name> <name><surname>Ma</surname><given-names>Y.</given-names></name> <name><surname>Wang</surname><given-names>X.</given-names></name> <name><surname>Wang</surname><given-names>X.</given-names></name> <name><surname>Li</surname><given-names>J.</given-names></name> <name><surname>Tian</surname><given-names>Y.</given-names></name> <etal/></person-group>. (<year>2022</year>). <article-title>Producing high-quality cultivation substrates for cucumber production by in-situ composting of corn straw blocks amended with biochar and earthworm casts</article-title>. <source>Waste Manag.</source> <volume>139</volume>, <fpage>179</fpage>&#x2013;<lpage>189</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.wasman.2021.12.010</pub-id>, <pub-id pub-id-type="pmid">34973573</pub-id></mixed-citation></ref>
<ref id="ref57"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Zhou</surname><given-names>L. M.</given-names></name> <name><surname>Jin</surname><given-names>S. L.</given-names></name> <name><surname>Liu</surname><given-names>C. A.</given-names></name> <name><surname>Xiong</surname><given-names>Y. C.</given-names></name> <name><surname>Si</surname><given-names>J. T.</given-names></name> <name><surname>Li</surname><given-names>X. G.</given-names></name> <etal/></person-group>. (<year>2012</year>). <article-title>Ridge-furrow and plastic-mulching tillage enhances maize&#x2013;soil interactions: opportunities and challenges in a semiarid agroecosystem</article-title>. <source>Field Crop Res.</source> <volume>126</volume>, <fpage>181</fpage>&#x2013;<lpage>188</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.fcr.2011.10.010</pub-id></mixed-citation></ref>
<ref id="ref58"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Zhu</surname><given-names>X.</given-names></name> <name><surname>Peng</surname><given-names>C.</given-names></name> <name><surname>Si</surname><given-names>G.</given-names></name> <name><surname>Sha</surname><given-names>A.</given-names></name> <name><surname>Yuan</surname><given-names>J.</given-names></name> <name><surname>Zhao</surname><given-names>S.</given-names></name> <etal/></person-group>. (<year>2022</year>). <article-title>Effects of straw returning on soil chemical properties and microbial community diversity under the rice-crayfish integrated system</article-title>. <source>Sustainability</source> <volume>14</volume>:<fpage>13539</fpage>. doi: <pub-id pub-id-type="doi">10.3390/su142013539</pub-id></mixed-citation></ref>
<ref id="ref59"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Zribi</surname><given-names>W.</given-names></name> <name><surname>Arag&#x00FC;&#x00E9;s</surname><given-names>R.</given-names></name> <name><surname>Medina</surname><given-names>E.</given-names></name> <name><surname>Faci</surname><given-names>J. M.</given-names></name></person-group> (<year>2015</year>). <article-title>Efficiency of inorganic and organic mulching materials for soil evaporation control</article-title>. <source>Soil Tillage Res.</source> <volume>148</volume>, <fpage>40</fpage>&#x2013;<lpage>45</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.still.2014.12.003</pub-id></mixed-citation></ref>
</ref-list>
<fn-group>
<fn fn-type="custom" custom-type="edited-by" id="fn0001">
<p>Edited by: <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/1862455/overview">Brahim Bouizgarne</ext-link>, Ibn Zohr University, Morocco</p>
</fn>
<fn fn-type="custom" custom-type="reviewed-by" id="fn0002">
<p>Reviewed by: <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/2576443/overview">Azim Khalid</ext-link>, National Institute for Agricultural Research, Morocco</p>
<p><ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/2707451/overview">Jamal Hallam</ext-link>, National Institute for Agricultural Research, Morocco</p>
</fn>
</fn-group>
</back>
</article>