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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Plant Sci.</journal-id>
<journal-title>Frontiers in Plant Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Plant Sci.</abbrev-journal-title>
<issn pub-type="epub">1664-462X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fpls.2025.1659478</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Plant Science</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Multi-environment meta-analysis reveals the mechanism of action of potassium-solubilizing microorganisms on crop yields</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Fan</surname>
<given-names>Xing</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/3121856/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Zhu</surname>
<given-names>Yuanpeng</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Jia</surname>
<given-names>Yan</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Du</surname>
<given-names>Peishen</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Wang</surname>
<given-names>Weini</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/funding-acquisition/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Liu</surname>
<given-names>Junmei</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/funding-acquisition/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Lv</surname>
<given-names>Zhijun</given-names>
</name>
<xref ref-type="aff" rid="aff5">
<sup>5</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/funding-acquisition/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Liu</surname>
<given-names>Ronghao</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="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2224675/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Li</surname>
<given-names>Xiaobin</given-names>
</name>
<xref ref-type="aff" rid="aff6">
<sup>6</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
</contrib>
</contrib-group>
<aff id="aff1">
<sup>1</sup>
<institution>College of Water Resources Science and Engineering, Taiyuan University of Technology</institution>, <addr-line>Taiyuan</addr-line>,&#xa0;<country>China</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>Shanxi Key Laboratory of Collaborative Utilization of River Basin Water Resources</institution>, <addr-line>Taiyuan</addr-line>,&#xa0;<country>China</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>Anhui Provincial Academy of Eco-Environmental Science Research</institution>, <addr-line>Hefei</addr-line>,&#xa0;<country>China</country>
</aff>
<aff id="aff4">
<sup>4</sup>
<institution>Ordos Agriculture and Animal Husbandry Ecology and Resource Protection Center</institution>, <addr-line>Ordos, Inner Mengulia</addr-line>,&#xa0;<country>China</country>
</aff>
<aff id="aff5">
<sup>5</sup>
<institution>Agricultural and Animal Husbandry Technology Extension Center, Dalat Banner</institution>, <addr-line>Ordos, Inner Mengulia</addr-line>,&#xa0;<country>China</country>
</aff>
<aff id="aff6">
<sup>6</sup>
<institution>Institute of Agricultural Resources and Regional Planning, Chinese Academy of Agricultural Sciences</institution>, <addr-line>Beijing</addr-line>,&#xa0;<country>China</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/177320/overview">M. J. I. Shohag</ext-link>, University of Florida, United States</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/451184/overview">Shafaqat Ali</ext-link>, Government College University, Pakistan</p>
<p>
<ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/2190554/overview">Abdoulaye Soumare</ext-link>, Ziguinchor University, Senegal</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Ronghao Liu, <email xlink:href="mailto:liuronghao@tyut.edu.cn">liuronghao@tyut.edu.cn</email>; Xiaobin Li, <email xlink:href="mailto:lixiaobin01@caas.cn">lixiaobin01@caas.cn</email>
</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>03</day>
<month>11</month>
<year>2025</year>
</pub-date>
<pub-date pub-type="collection">
<year>2025</year>
</pub-date>
<volume>16</volume>
<elocation-id>1659478</elocation-id>
<history>
<date date-type="received">
<day>04</day>
<month>07</month>
<year>2025</year>
</date>
<date date-type="accepted">
<day>20</day>
<month>10</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2025 Fan, Zhu, Jia, Du, Wang, Liu, Lv, Liu and Li.</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>Fan, Zhu, Jia, Du, Wang, Liu, Lv, Liu and Li</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>The availability of soil potassium plays a critical role in yield increases. Potassium-solubilizing microorganisms (KSM) offer a promising biological solution to improve potassium availability, but their efficacy across diverse global environments remains uncertain. Through a global meta-analysis of 102 studies (846 paired observations), we systematically evaluated the effects of KSM application on crop yields across five key dimensions: microorganism types, soil factors, crop classifications, field management, and stress types. KSM inoculation significantly increased soil available potassium (+28.9%), crop yield (+23.4%), and key growth indices, such as root length (+29.50%) and leaf area (+44.7%). This study identified <italic>Aspergillus</italic> spp. as the most suitable microorganism, and revealed that KSM efficacy is highly dependent on context: yield responses were greatest in clay loam soils, vegetable crops, and greenhouse conditions. Structural equation modeling indicated that microbial abundance, climate, soil available potassium, and plant growth (root length and leaf area) are key direct and indirect drivers of yield enhancement. The results indicate that the application of KSM is an effective strategy to increase crop yields in various environments. By identifying the optimal conditions for KSM application, the identification of optimal application parameters, derived from cross-study analysis, provides a robust strategy for leveraging microbial communities to boost soil potassium availability and nutrient efficiency, thereby contributing to the transition toward more sustainable and climate-resilient agriculture.</p>
</abstract>
<kwd-group>
<kwd>potassium-solubilizing microorganisms</kwd>
<kwd>soil available potassium</kwd>
<kwd>crop yield</kwd>
<kwd>meta-analysis</kwd>
<kwd>sustainable agriculture</kwd>
</kwd-group>
<counts>
<fig-count count="7"/>
<table-count count="1"/>
<equation-count count="6"/>
<ref-count count="44"/>
<page-count count="12"/>
<word-count count="4946"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Plant Nutrition</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>Potassium (K) is an essential plant nutrient and the most abundant cation in plant cells, playing a critical role in a myriad of physiological and metabolic processes, including enzyme activation, osmotic regulation, and photosynthesis (<xref ref-type="bibr" rid="B33">Sheng, 2005</xref>; <xref ref-type="bibr" rid="B30">Sattar et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B20">Li et&#xa0;al., 2023b</xref>). Consequently, potassium availability directly governs photosynthetic efficiency, crop stress resistance, growth, and ultimately yield and quality (<xref ref-type="bibr" rid="B42">Yang et&#xa0;al., 2023</xref>; <xref ref-type="bibr" rid="B40">Wang et&#xa0;al., 2024</xref>; <xref ref-type="bibr" rid="B43">Zhao et&#xa0;al., 2024</xref>). However, the natural soil potassium pool often lacks sufficient capacity to convert insoluble K into plant-available forms, frequently failing to meet crop demand (<xref ref-type="bibr" rid="B41">Xu et&#xa0;al., 2025</xref>). This limitation represents a fundamental constraint on agricultural productivity and sustainability, contributing to inefficient fertilizer use and environmental imbalances (<xref ref-type="bibr" rid="B43">Zhao et&#xa0;al., 2024</xref>; <xref ref-type="bibr" rid="B21">Liu et&#xa0;al., 2025</xref>).</p>
<p>Addressing this challenge requires a shift from purely chemical solutions towards biological strategies that harness soil ecosystem processes. Potassium-solubilizing microorganisms (KSM) are a key functional group within the soil microbiome that can enhance mineral weathering and mobilize fixed potassium pools through biological mineralization (<xref ref-type="bibr" rid="B30">Sattar et&#xa0;al., 2019</xref>). Their mechanisms (including acid production, phytohormone secretion, and extracellular enzyme release) not only improve potassium availability but also enhance soil health and plant resilience (<xref ref-type="bibr" rid="B24">Meena et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B35">Sun et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B4">Chen et&#xa0;al., 2022</xref>). This positions KSM application at the intersection of microbial ecology and plant nutrition, offering a pathway to more sustainable agricultural intensification by working with, rather than against, soil biological processes (<xref ref-type="bibr" rid="B32">Shen et&#xa0;al., 2016</xref>).</p>
<p>Numerous targeted studies have demonstrated the efficacy of specific KSM strains (e.g., <italic>Bacillus</italic>, <italic>Pseudomonas</italic>) in increasing potassium solubility and crop yields under controlled or local conditions (<xref ref-type="bibr" rid="B17">Khanghahi et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B29">Sarikhani et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B26">Nawaz et&#xa0;al., 2023</xref>). However, the transition from promising microbe-level effects to reliable field-scale outcomes remains a major hurdle in microbial ecology applied to agriculture. A critical gap exists between demonstrating efficacy in discrete settings and predicting effectiveness across the heterogeneous landscapes of global agriculture. The intricate relationships between KSM functionality, environmental context (e.g., soil type, stress type), and field management are poorly quantified. Consequently, it is unclear whether the benefits of KSM are generalizable or contingent on specific, and often unknown, ecological interactions and agronomic conditions. This uncertainty hinders the development of predictive models and evidence-based recommendations for integrating KSM into sustainable farming systems.</p>
<p>To bridge this gap between microbial ecology and agricultural practice, we conducted a comprehensive global meta-analysis of studies from 2015-2024. By synthesizing evidence across five key dimensions: microbial species, soil types, crop types, field management, and stress types (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>). We aim to: (i) quantify the overarching impact of KSM on soil potassium dynamics and plant performance; (ii) identify the contextual factors that most strongly regulate KSM efficacy; and (iii) elucidate the complex pathways through which KSM influence crop yield, integrating microbial, plant, and environmental variables. Our work provides a mechanistic, evidence-based framework for understanding KSM ecology in agricultural systems, ultimately guiding the development of targeted microbial strategies for enhancing potassium use efficiency and advancing sustainable crop production.</p>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Subgroup classification table for the forest plots.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" rowspan="2" align="center">First-order factor</th>
<th valign="middle" rowspan="2" align="center">Secondary factor</th>
<th valign="middle" colspan="5" align="center">Clusters</th>
</tr>
<tr>
<th valign="middle" align="center">1</th>
<th valign="middle" align="center">2</th>
<th valign="middle" align="center">3</th>
<th valign="middle" align="center">4</th>
<th valign="middle" align="center">5</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="center">Microorganism</td>
<td valign="middle" align="center">Potassium-solubilizing microorganisms</td>
<td valign="middle" align="center">Bacillus</td>
<td valign="middle" align="center">Enterobacteria</td>
<td valign="middle" align="center">Aspergillus</td>
<td valign="middle" align="center">Pseudomonas</td>
<td valign="middle" align="center">Rhizobium</td>
</tr>
<tr>
<td valign="middle" rowspan="2" align="center">Soil factor</td>
<td valign="middle" align="center">Soil texture</td>
<td valign="middle" align="center">Sandy loam</td>
<td valign="middle" align="center">Loam</td>
<td valign="middle" align="center">Clay loam</td>
<td valign="middle" align="center">&#x2013;</td>
<td valign="middle" align="center">&#x2013;</td>
</tr>
<tr>
<td valign="middle" align="center">Soil pH</td>
<td valign="middle" align="center">&gt; 8</td>
<td valign="middle" align="center">&gt; 6&#x2013;8</td>
<td valign="middle" align="center">&lt; 6</td>
<td valign="middle" align="center">&#x2013;</td>
<td valign="middle" align="center">&#x2013;</td>
</tr>
<tr>
<td valign="middle" rowspan="2" align="center">Crop classification</td>
<td valign="middle" align="center">Crop category</td>
<td valign="middle" align="center">Vegetable crop</td>
<td valign="middle" align="center">Food crop</td>
<td valign="middle" align="center">Fruit tree</td>
<td valign="middle" align="center">&#x2013;</td>
<td valign="middle" align="center">&#x2013;</td>
</tr>
<tr>
<td valign="middle" align="center">Crop family</td>
<td valign="middle" align="center">Poaceae</td>
<td valign="middle" align="center">Legume</td>
<td valign="middle" align="center">Solanaceae</td>
<td valign="middle" align="center">Cruciferous</td>
<td valign="middle" align="center">Anacardiaceae</td>
</tr>
<tr>
<td valign="middle" align="center">Field management</td>
<td valign="middle" align="center">Planting method</td>
<td valign="middle" align="center">Greenhouse experiment</td>
<td valign="middle" align="center">Field experiment</td>
<td valign="middle" align="center">&#x2013;</td>
<td valign="middle" align="center">&#x2013;</td>
<td valign="middle" align="center">&#x2013;</td>
</tr>
<tr>
<td valign="middle" rowspan="2" align="center">Stress type</td>
<td valign="middle" align="center">Soil water potential</td>
<td valign="middle" align="center">-25~&#x2013;10 kPa</td>
<td valign="middle" align="center">-50~&#x2013;25 kPa</td>
<td valign="middle" align="center">-80~&#x2013;50 kPa</td>
<td valign="middle" align="center">&#x2013;</td>
<td valign="middle" align="center">&#x2013;</td>
</tr>
<tr>
<td valign="middle" align="center">Degree of soil salinization</td>
<td valign="middle" align="center">4.0&#x2013;8.0 dSm<sup>&#x2013;1</sup>
</td>
<td valign="middle" align="center">8.0&#x2013;16.0 dSm<sup>&#x2013;1</sup>
</td>
<td valign="middle" align="center">&gt; 16.0 dSm<sup>&#x2013;1</sup>
</td>
<td valign="middle" align="center">&#x2013;</td>
<td valign="middle" align="center">&#x2013;</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="s2" sec-type="materials|methods">
<label>2</label>
<title>Materials and methods</title>
<sec id="s2_1">
<label>2.1</label>
<title>Data retrieval strategies and filtering criteria</title>
<p>The data for the meta-analysis were retrieved from Web of Science (WOS) and the China National Knowledge Infrastructure (CNKI), and the keywords searched included &#x201c;potassium-solubilizing bacteria&#x201d;, &#x201c;yield&#x201d;, &#x201c;rhizosphere potassium-solubilizing bacteria&#x201d;, &#x201c;microorganisms&#x201d;, &#x201c;enzyme activity&#x201d; and &#x201c;potassium solubilization&#x201d;. For the meta-analysis, 102 relevant published studies were collated, including 95 English and 7 Chinese studies (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>), containing 846 pairs of paired observations(Among them, 806 pairs of data are from English databases, and 40 pairs of data are from Chinese databases). The selected studies had to satisfy the following requirements: (1) The retrieved variables included the mean, standard deviation (or standard error), and repeat group. (2) All the control and treatment groups belonged to the same ecosystem and experienced the same environmental and growth conditions. (3) Data from only the control and treatment groups inoculated with potassium-solubilizing microorganisms were analyzed in multiple controlled studies with experimental inoculation treatments. (4) The extracted literature was limited to articles published from 2015 to 2025.</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Map of study sites included in the meta-analysis.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1659478-g001.tif">
<alt-text content-type="machine-generated">World map showing experimental sites marked by blue triangles. Sites are distributed globally, with clusters in East Asia, Europe, and scattered points across North and South America. Latitude and longitude lines provide geographic reference. A scale bar indicates distances up to eleven thousand kilometers.</alt-text>
</graphic>
</fig>
</sec>
<sec id="s2_2">
<label>2.2</label>
<title>Data collection and classification</title>
<p>We also recorded climatic characteristics such as the mean annual temperature (MAT) and mean annual precipitation (MAP), collecting climatic data from the NASA Surface Meteorology and Solar Energy Location site (<ext-link ext-link-type="uri" xlink:href="https://power.larc.nasa.gov/">https://power.larc.nasa.gov/</ext-link>) on the basis of the latitude and longitude of the study site, if not reported in the original study.</p>
<p>Data from all published literature tables were extracted manually, whereas data from figures were extracted via the Get Data (<ext-link ext-link-type="uri" xlink:href="https://getdata.sourceforge.net/download.html">https://getdata.sourceforge.net/download.html</ext-link>) tool.</p>
</sec>
<sec id="s2_3">
<label>2.3</label>
<title>Meta-data analysis</title>
<p>We used Meta-Win 2.0 software (<ext-link ext-link-type="uri" xlink:href="https://www.metawinsoft.com/">https://www.metawinsoft.com/</ext-link>) to analyze the data for random effects. The effect value was calculated via the natural logarithm of the response ratio (lnRR) in <xref ref-type="disp-formula" rid="eq2">Equation (2)</xref> (<xref ref-type="bibr" rid="B22">Luo et&#xa0;al., 2018</xref>), which was then converted to upper and lower 95% confidence intervals (CI) in <xref ref-type="disp-formula" rid="eq4">Equation (4)</xref> to evaluate the impact of the KSM on crop yield, soil physicochemical parameters, and crop growth indicators. In several studies, the standard error (SE) in <xref ref-type="disp-formula" rid="eq1">Equation (1)</xref> had to be translated to the standard deviation (SD) in <xref ref-type="disp-formula" rid="eq1">Equation (1)</xref>, which was calculated as (<xref ref-type="bibr" rid="B15">Jian et&#xa0;al., 2016</xref>):</p>
<disp-formula id="eq1">
<label>(1)</label>
<mml:math display="block" id="M1">
<mml:mrow>
<mml:mtext>SD</mml:mtext>
<mml:mo>=</mml:mo>
<mml:mtext>SE</mml:mtext>
<mml:mo>&#xd7;</mml:mo>
<mml:msqrt>
<mml:mtext>n</mml:mtext>
</mml:msqrt>
</mml:mrow>
</mml:math>
</disp-formula>
<disp-formula id="eq2">
<label>(2)</label>
<mml:math display="block" id="M2">
<mml:mrow>
<mml:mtext>lnRR&#xa0;=ln</mml:mtext>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mrow>
<mml:mfrac>
<mml:mrow>
<mml:mover accent="true">
<mml:mrow>
<mml:msub>
<mml:mi>x</mml:mi>
<mml:mi>t</mml:mi>
</mml:msub>
</mml:mrow>
<mml:mo stretchy="true">&#xaf;</mml:mo>
</mml:mover>
</mml:mrow>
<mml:mrow>
<mml:mover accent="true">
<mml:mrow>
<mml:msub>
<mml:mi>x</mml:mi>
<mml:mi>c</mml:mi>
</mml:msub>
</mml:mrow>
<mml:mo stretchy="true">&#xaf;</mml:mo>
</mml:mover>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:mrow>
</mml:math>
</disp-formula>
<p>The variance of the effect value is as follows:</p>
<disp-formula id="eq3">
<label>(3)</label>
<mml:math display="block" id="M3">
<mml:mrow>
<mml:mi>var</mml:mi>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi>l</mml:mi>
<mml:mi>n</mml:mi>
<mml:mi>R</mml:mi>
<mml:mi>R</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
<mml:mo>=</mml:mo>
<mml:mfrac>
<mml:mrow>
<mml:msubsup>
<mml:mi>S</mml:mi>
<mml:mi>t</mml:mi>
<mml:mn>2</mml:mn>
</mml:msubsup>
</mml:mrow>
<mml:mrow>
<mml:msub>
<mml:mi>n</mml:mi>
<mml:mi>t</mml:mi>
</mml:msub>
<mml:msubsup>
<mml:mover accent="true">
<mml:mi>X</mml:mi>
<mml:mo>&#xaf;</mml:mo>
</mml:mover>
<mml:mi>t</mml:mi>
<mml:mn>2</mml:mn>
</mml:msubsup>
</mml:mrow>
</mml:mfrac>
<mml:mo>+</mml:mo>
<mml:mfrac>
<mml:mrow>
<mml:msubsup>
<mml:mi>S</mml:mi>
<mml:mi>c</mml:mi>
<mml:mn>2</mml:mn>
</mml:msubsup>
</mml:mrow>
<mml:mrow>
<mml:msub>
<mml:mi>n</mml:mi>
<mml:mi>c</mml:mi>
</mml:msub>
<mml:msubsup>
<mml:mover accent="true">
<mml:mi>X</mml:mi>
<mml:mo>&#xaf;</mml:mo>
</mml:mover>
<mml:mi>c</mml:mi>
<mml:mn>2</mml:mn>
</mml:msubsup>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
</mml:math>
</disp-formula>
<p>where <inline-formula>
<mml:math display="inline" id="im1">
<mml:mrow>
<mml:msub>
<mml:mover accent="true">
<mml:mi>X</mml:mi>
<mml:mo>&#xaf;</mml:mo>
</mml:mover>
<mml:mi>t</mml:mi>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> is the mean of the variable in the treatment group and where <inline-formula>
<mml:math display="inline" id="im2">
<mml:mrow>
<mml:msub>
<mml:mover accent="true">
<mml:mi>X</mml:mi>
<mml:mo>&#xaf;</mml:mo>
</mml:mover>
<mml:mi>c</mml:mi>
</mml:msub>
</mml:mrow>
</mml:math>
</inline-formula> is the mean of the variable in the control group in <xref ref-type="disp-formula" rid="eq3">Equation (3)</xref>. S<sub>t</sub> is the standard deviation of the variable in the treatment group; S<sub>c</sub> is the standard deviation of the variable in the control group; n is the sample size; and n<sub>t</sub> and n<sub>c</sub> are the sample sizes of the treatment and control groups, respectively in <xref ref-type="disp-formula" rid="eq3">Equation (3)</xref>. We converted lnRR and its corresponding confidence intervals to the corresponding percentage change.</p>
<disp-formula id="eq4">
<label>(4)</label>
<mml:math display="block" id="M4">
<mml:mrow>
<mml:mi>C</mml:mi>
<mml:mi>I</mml:mi>
<mml:mo>=</mml:mo>
<mml:mo stretchy="false">(</mml:mo>
<mml:msup>
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<mml:mrow>
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<mml:mi>R</mml:mi>
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<mml:mn>100</mml:mn>
<mml:mo>%</mml:mo>
</mml:mrow>
</mml:math>
</disp-formula>
</sec>
<sec id="s2_4">
<label>2.4</label>
<title>Heterogeneity test and publication bias test</title>
<p>The I<sup>2</sup> statistic was used to assess heterogeneity, and the test value Q was calculated as follows in <xref ref-type="disp-formula" rid="eq5">Equation (5)</xref> and <xref ref-type="disp-formula" rid="eq6">(6)</xref> (<xref ref-type="bibr" rid="B12">Huo et&#xa0;al., 2017</xref>):</p>
<disp-formula id="eq5">
<label>(5)</label>
<mml:math display="block" id="M5">
<mml:mrow>
<mml:mtext>Q</mml:mtext>
<mml:mo>=</mml:mo>
<mml:msubsup>
<mml:mo>&#x2211;</mml:mo>
<mml:mrow>
<mml:mtext>i</mml:mtext>
<mml:mo>=</mml:mo>
<mml:mn>1</mml:mn>
</mml:mrow>
<mml:mtext>n</mml:mtext>
</mml:msubsup>
<mml:msub>
<mml:mtext>W</mml:mtext>
<mml:mtext>i</mml:mtext>
</mml:msub>
<mml:msup>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:msubsup>
<mml:mi>E</mml:mi>
<mml:mi>i</mml:mi>
<mml:mn>2</mml:mn>
</mml:msubsup>
<mml:mo>&#x2212;</mml:mo>
<mml:msup>
<mml:mtext>E</mml:mtext>
<mml:mn>2</mml:mn>
</mml:msup>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
<mml:mn>2</mml:mn>
</mml:msup>
</mml:mrow>
</mml:math>
</disp-formula>
<disp-formula id="eq6">
<label>(6)</label>
<mml:math display="block" id="M6">
<mml:mrow>
<mml:msup>
<mml:mi>I</mml:mi>
<mml:mn>2</mml:mn>
</mml:msup>
<mml:mo>=</mml:mo>
<mml:mo stretchy="false">[</mml:mo>
<mml:mi>Q</mml:mi>
<mml:mo>&#x2212;</mml:mo>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi>n</mml:mi>
<mml:mo>&#x2212;</mml:mo>
<mml:mn>1</mml:mn>
<mml:mo stretchy="false">)</mml:mo>
<mml:mo stretchy="false">]</mml:mo>
<mml:mo stretchy="false">/</mml:mo>
<mml:mi>Q</mml:mi>
</mml:mrow>
</mml:math>
</disp-formula>
<p>where w<sub>i</sub> is the weight of group i of data, n is the number of effect sizes, E<sub>i</sub> is the effect size of the ith group, and E is the average of all data effect sizes in <xref ref-type="disp-formula" rid="eq5">Equation (5)</xref>. Calculations were made using the random effect model when the heterogeneity test result I<sup>2</sup> was greater than 50% and the fixed effect model when it was less than 50%. The Rosenthal loss of safety coefficient method was used for the publication bias test. N&gt;5 m+10 (N is the loss of safety coefficient, and m is the sample size) indicates no publication bias. We used AMOS software (IBM SPSS AMOS 20.0.0) to untangle the indirect and direct effects of climate (MAT and MAP), microbial count, soil available potassium content, crop growth indicators, and soil enzyme activities on crop yields (<xref ref-type="bibr" rid="B6">Eisenhauer et&#xa0;al., 2015</xref>) via structural equation modeling (SEM). Before modeling, all the data were normalized (ensuring that the data followed a standard normal distribution with a mean value of 0 and standard deviation of 1), and an <italic>a priori</italic> model was designed based on the known effects and relationships among the drivers that had a significant impact on crop yields in our previous analyses. SEM was used instead of multiple regressions since directions can be assigned to several relationships, resulting in multiple explanatory and response variables in one model. Furthermore, the structure of such a model can reveal whether a significant bivariate relationship results from a significant relationship between two given variables and a third variable. Despite using fail-safe N to assess publication bias, our results may still be affected by the underreporting of studies with non-significant or negative findings. While subgroup analysis explored substantial heterogeneity (indicated by I&#xb2;), residual heterogeneity could remain due to unmeasured factors like experimental duration, KSM strain potency, and unrecorded soil microbial communities, which may limit the generalizability of pooled effects (<xref ref-type="bibr" rid="B16">Jin et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B23">Mathur and VanderWeele, 2022</xref>).</p>
</sec>
</sec>
<sec id="s3" sec-type="results">
<label>3</label>
<title>Results</title>
<sec id="s3_1">
<label>3.1</label>
<title>Effect of KSM inoculation on crop yield</title>
<p>The calculation of the effect size on yield revealed that KSM inoculation improved overall crop yield by 23.43% compared with the control, confirming its potential as a yield-enhancing agricultural practice (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>). Note: The four KSM genera analyzed (<italic>Aspergillus</italic>, <italic>Pseudomonas</italic>, <italic>Bacillus</italic>, <italic>Enterobacter</italic>) were selected based on two criteria: (1) their high frequency of reporting in existing KSM-related studies (accounting for &gt;70% of eligible literature included in this meta-analysis); (2) their well-documented potassium-solubilizing mechanisms and agricultural application records, ensuring data robustness and practical relevance. Among these genera, <italic>Aspergillus</italic> (+36.27%) and <italic>Pseudomonas</italic> (+32.86%) were consistently demonstrated to outperform <italic>Bacillus</italic> and <italic>Enterobacter</italic> in boosting crop yields (P&lt;0.05).</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Effect value of KSM inoculation over uninoculated KSM on crop yield (the significant signal means *P&lt;0.05, **P&lt;0.01, ***P&lt;0.001).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1659478-g002.tif">
<alt-text content-type="machine-generated">Forest plot showing weighted response ratio (RR-) for various factors affecting yield. Categories include microbial species, soil textures, pH levels, crop types, crop families, planting methods, soil water potential, and soil electrical conductivity. Each factor&#x2019;s RR- is represented by a dot with confidence intervals, and the number of studies is indicated in parentheses. Significant results are marked by asterisks. The vertical dashed line denotes zero effect.</alt-text>
</graphic>
</fig>
<p>Soil texture emerged as a key modulating factor: clay loam soil (+38.82%) showed a significantly stronger yield response to KSM than sandy loam and loam (P&lt;0.05), while soil pH had no substantial impact. Crop type and family also drove marked differences: vegetable crops (+34.25%) exhibited the greatest yield increase relative to food crops and fruit trees (P&lt;0.05), with cruciferous (+42.74%) and leguminous (+40.08%) crops outperforming Poaceae and Anacardiaceae (P&lt;0.05). Additionally, greenhouse experiments (+34.17%) yielded better results than field trials (P&lt;0.05), whereas soil water potential and EC values did not induce significant variations in yield response.</p>
</sec>
<sec id="s3_2">
<label>3.2</label>
<title>Effect of KSM inoculation on soil physicochemical properties</title>
<sec id="s3_2_1">
<label>3.2.1</label>
<title>Effect of KSM inoculation on soil nutrients and pH</title>
<p>Compared with the control, KSM inoculation significantly modified soil potassium status and pH, with soil total potassium and available potassium increasing by 18.28% and 28.91%, respectively, and soil pH decreasing by 18.10% (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Figures S2A, B</bold>
</xref>).</p>
<p>Notably, the efficacy of KSM in boosting soil available potassium varied markedly by biological and environmental factors: <italic>Enterobacter</italic> (+39.22%) showed a significantly stronger capacity to mobilize available potassium than <italic>Bacillus</italic> and <italic>Pseudomonas</italic> (P&lt;0.05), while loam soil (+32.25%) outperformed clay loam and sandy loam in this regard (P&lt;0.05). Crop-related factors also drove substantial differences: vegetable crops (+50.4%) and Poaceae (+32.71%) exerted the greatest positive effects on soil potassium availability among crop types and families, respectively (P&lt;0.05). In contrast, soil pH had no substantial impact on KSM-mediated available potassium augmentation, indicating that KSM can stably improve potassium availability across diverse soil acid-base conditions.</p>
<p>For soil pH regulation, all tested KSM genera induced negative effects, though with no significant differences between them. Soil texture and initial pH modulated this effect: loam (&#x2013;18.32%) and clay loam (&#x2013;25.06%) showed stronger pH reduction than sandy loam, and KSM inoculation at pH&gt;8 (&#x2013;25.38%) had a significantly greater acidifying effect than at pH 6&#x2013;8 (P&lt;0.05), suggesting that KSM are more effective at lowering pH in alkaline soils.</p>
</sec>
<sec id="s3_2_2">
<label>3.2.2</label>
<title>Effect of KSM inoculation on soil enzyme activity</title>
<p>KSM inoculation significantly enhanced the activities of both antioxidant-related and metabolism-related enzymes, with peroxidase (+41.0%) and cellulase (+147.3%) showing the highest enhancement rates, respectively (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Figure S2A, C</bold>
</xref>).</p>
<p>For antioxidant-related enzymes, superoxide dismutase activity trended positive in sandy loam and loam but decreased in clay loam, while peroxidase was beneficial only in loam soil. In contrast, catalase activity showed clear strain and texture specificity: <italic>Pseudomonas</italic> (+50.92%) was significantly more effective than <italic>Bacillus</italic> (P&lt;0.05), and clay loam/loam supported positive responses whereas sandy loam showed a negative effect. Consistently across all antioxidant enzymes, KSM promoted activity at soil pH&lt;6 but suppressed it at pH&gt;8 relative to the control, highlighting soil pH as a key regulator of KSM-induced antioxidant defense.</p>
<p>Metabolism-related enzymes showed universal enhancement, with protease (+58.9%), chitinase (+62.7%), and cellulase (+147.3%) exhibiting the strongest activity increases, indicating that KSM play a critical role in accelerating soil organic matter decomposition and nutrient cycling. Sucrase showed the weakest response (+13.6%), suggesting that KSM have enzyme-specific regulatory effects on soil metabolic processes.</p>
</sec>
</sec>
<sec id="s3_3">
<label>3.3</label>
<title>Effect of KSM inoculation on crop growth</title>
<sec id="s3_3_1">
<label>3.3.1</label>
<title>Effect of KSM inoculation on crop potassium content</title>
<p>KSM inoculation significantly enhanced potassium uptake in crops, with root K concentration and shoot K concentration both showing marked increases relative to the control (<xref ref-type="fig" rid="f3">
<bold>Figures&#xa0;3A, B</bold>
</xref>).</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Effect of different conditions on K concentrations in roots <bold>(A)</bold> and in shoots <bold>(B)</bold> under inoculation with KSM (the significant signal means *P&lt;0.05, **P&lt;0.01, ***P&lt;0.001).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1659478-g003.tif">
<alt-text content-type="machine-generated">Two panels, A and B, depict weighted response ratios (RR-) related to potassium (K) concentrations in roots and shoots against various factors. Panel A focuses on root K concentrations, showing data for bacteria, soil texture, crop categories, and soil characteristics. Panel B highlights shoot K concentrations with similar variables, adding more categories like planting methods. Each factor has associated RR values and confidence intervals, indicating variability and significance levels. Both panels are organized by factors, such as soil type and crop family, with total, specific bacterial, and soil conditions illustrated.</alt-text>
</graphic>
</fig>
<p>For root K concentration, <italic>Aspergillus</italic> (+79.64%) was the most effective genus, outperforming <italic>Bacillus</italic> and <italic>Pseudomonas</italic> (P&lt;0.05). Notably, the effect values for root K concentration were consistently positive across soil textures, crop types, crop families, and soil ECe conditions, with no significant variations between subgroups&#x2014;indicating that KSM promotes root potassium accumulation stably across diverse contexts.</p>
<p>Shoot K concentration was increased by 31.11% overall, with <italic>Aspergillus</italic> (+77.65%) again outperforming <italic>Enterobacter</italic>, <italic>Bacillus</italic>, and <italic>Pseudomonas</italic> (P&lt;0.05). Soil texture significantly modulated this effect: clay loam (+55.99%) and sandy loam (+53.71%) were more effective than loam (P&lt;0.05). By crop category, vegetable crops (+60.38%) showed a stronger response than food crops and fruit trees (P&lt;0.05), and Solanaceae (+60.67%) outperformed other crop families (P&lt;0.05). No significant differences in shoot K concentration were detected under various soil ECe conditions.</p>
</sec>
<sec id="s3_3_2">
<label>3.3.2</label>
<title>Effect of KSM inoculation on growth indices</title>
<p>Compared with the control, KSM significantly increased all measured crop growth indices, with dry root weight (+59.21%) and total chlorophyll content (+56.06%) showing the most dramatic improvements, followed by leaf area (+44.67%) and root fresh weight (+40.79%) (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Figure S3A</bold>
</xref>).</p>
<p>For root length (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4A</bold>
</xref>), KSM exerted positive effects across all subgroups, with clay loam soil (+52.71%) and field experiments (+57.20%) standing out as the most favorable conditions (P&lt;0.05). Notably, plant height responded strongly to <italic>Aspergillus</italic> inoculation (+54.33%), which was significantly more effective than other genera (P&lt;0.05); sandy loam soil (+36.71%) and vegetable crops (+43.30%), particularly solanaceae (+47.11%), also showed enhanced plant height under KSM treatment (P&lt;0.05). In contrast, soil pH had no significant influence on plant height. Greenhouse experiments (+49.86%) outperformed field trials in promoting plant height (P&lt;0.05), and the optimal soil water potential range was &#x2013;80 to &#x2013;50 kPa (+55.64%, P&lt;0.05).</p>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Effect of different conditions on root length <bold>(A)</bold>, leaf area <bold>(B)</bold> under inoculation with KSM (the significant signal means *P&lt;0.05, **P&lt;0.01, ***P&lt;0.001).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1659478-g004.tif">
<alt-text content-type="machine-generated">Forest plot comparing root length and leaf area with various factors such as microorganisms, soil texture, soil pH, crop category, crop family, planting method, soil water potential, and soil electrical conductivity (ECe). Panel A focuses on root length, while Panel B focuses on leaf area. Each point is a weighted response ratio (RR-) with confidence intervals indicating variability. Dotted lines separate categories and factors, with numbers indicating sample sizes.</alt-text>
</graphic>
</fig>
<p>Leaf area was most notably increased by <italic>Aspergillus</italic> (+76.00%, P&lt;0.05), with vegetable crops (+62.89%) and solanaceae (+63.37%) again showing stronger responses than other crop types/families (P&lt;0.05, <xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4B</bold>
</xref>). Greenhouse conditions (+71.24%) also facilitated a greater increase in leaf area compared to field settings (P&lt;0.05), while soil texture and pH had no significant differential effects.</p>
</sec>
</sec>
<sec id="s3_4">
<label>3.4</label>
<title>Effect of KSM inoculation on the K content in soil and crops in relation to environmental factors</title>
<p>A key chain emerged from the correlation analysis (<xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5</bold>
</xref>): soil available potassium was significantly and positively correlated with both crop shoot K concentrations (P&lt;0.05) and root K concentrations (P&lt;0.05), while root K concentrations further positively linked to shoot K concentrations (P&lt;0.05). Critically, both soil available potassium and crop shoot K concentrations showed direct positive correlations with crop yield (P&lt;0.05), highlighting that KSM&#x2019;s yield-enhancing effect is likely mediated through improved soil K availability and subsequent crop K uptake.</p>
<fig id="f5" position="float">
<label>Figure&#xa0;5</label>
<caption>
<p>Distribution of the variable importance for <bold>(A)</bold> Yield, <bold>(B)</bold> Root length, <bold>(C)</bold> Plant height, <bold>(D)</bold> Leaf area, <bold>(E)</bold> Root K concentrations, and <bold>(F)</bold> Shoot K concentrations random forest models including the different Microbial count, climatic conditions, soil properties and Growth indicators as variable, the significant signal means *P&lt;0.05, **P&lt;0.01.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1659478-g005.tif">
<alt-text content-type="machine-generated">Bar chart panels labeled A to F show the increase in Mean Squared Error (MSE) percentage for different factors affecting yield, root length, plant height, leaf area, root K concentrations, and shoot K concentrations. Each panel lists factors on the y-axis with corresponding MSE increase percentages on the x-axis. Significant factors are marked with asterisks, with microbial count, soil available K, MAT, soil water potential, and soil pH frequently highlighted across the panels.</alt-text>
</graphic>
</fig>
</sec>
<sec id="s3_5">
<label>3.5</label>
<title>Relationships between effect sizes of inoculation with KSM on yield and other growth indicators</title>
<p>The results of the random forest analysis are presented in <xref ref-type="fig" rid="f6">
<bold>Figure&#xa0;6</bold>
</xref>. We assigned significance to factors influencing yield and other growth indicator changes by considering those with a significance level greater than 0.1. For yield, the key control factor is microbial count and soil available K. For root length, the key control factor is soil available K. For plant height, the key control factors are root length, soil available K, MAT, microbial count, soil pH, MAP, shoot K concentrations, and soil water potential. For leaf area, significant control factors included soil pH, microbial count, and plant height. For root K concentrations, significant control factors included microbial count, MAT, and soil available K. For shoot K concentrations, the significant control factors included soil available K and root K concentrations.</p>
<fig id="f6" position="float">
<label>Figure&#xa0;6</label>
<caption>
<p>Relationship between the effect size of soil available K and <bold>(A)</bold> yield, B) root K concentrations, <bold>(C)</bold> shoot K concentrations, <bold>(D)</bold> root length and <bold>(E)</bold> plant height; root K concentrations and <bold>(F)</bold> shoot K concentrations, <bold>(G)</bold> plant height and <bold>(H)</bold> leaf area; shoot K concentrations and <bold>(I)</bold> plant height, <bold>(G)</bold> leaf area; plant height and <bold>(K)</bold> yield. Reported statistical results from two-sided linear regressions. Gray areas represent 95%Cl.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1659478-g006.tif">
<alt-text content-type="machine-generated">Eleven scatter plots labeled A to K with trend lines and shaded confidence intervals. Each graph displays different relationships involving soil available potassium (K), root and shoot K concentrations, yield, plant height, and leaf area. The equations, R-squared, and P-values are presented on each plot, indicating statistical analyses of the relationships. Blue dots represent data points across all graphs.</alt-text>
</graphic>
</fig>
</sec>
<sec id="s3_6">
<label>3.6</label>
<title>Impact of microbial count, MAT, and MAP on crop yield following KSM inoculation</title>
<p>Structural equation modeling (SEM) was used to investigate how the various explanatory variables and their interactions influenced crop yield under KSM inoculation (<xref ref-type="fig" rid="f7">
<bold>Figure&#xa0;7</bold>
</xref>). The final structural equation model explained 19% of the overall variation in crop yield (&#x3c7;<sup>2</sup> = 5.09; P = 0.955, AIC = 113.09). The microbial count, MAP, MAT, soil available potassium, root K concentration, root length, leaf area, and oxidase were the most crucial factors affecting crop yield, both directly and indirectly. Overall, the microbial count indirectly affects crop yield by altering root length (path coefficient = 0.76), root K concentration (path coefficient = 0.93), and leaf area (path coefficient = 1.49). MAP indirectly affects crop yield by influencing root length (path coefficient = 0.81), root K concentration (path coefficient = 0.92), and leaf area (path coefficient = 1.69). The MAT had an indirect effect (path coefficient = 0.81), and the root K concentration (path coefficient = 0.92) and leaf area (path coefficient = 1.69) were affected. MAT also indirectly improved crop yield by influencing root K concentrations (path coefficient =&#x2013;0.21) and leaf area (path coefficient =&#x2013;0.15). Soil available potassium indirectly affected crop yield by influencing root length (path coefficient = 0.31) and root K concentrations (path coefficient = 0.29). The root K concentrations also indirectly influenced crop yield because of their impact on root length (path coefficient =&#x2013;0.32). Root length had a direct positive effect on crop yield (path coefficient = 0.21), and leaf area had a direct positive effect on crop yield (path coefficient = 0.24). Oxidase had a direct positive effect on crop yield (path coefficient = 0.14). These results confirm the potential of KSM to increase crop production through inoculation.</p>
<fig id="f7" position="float">
<label>Figure&#xa0;7</label>
<caption>
<p>Structural equation modeling (SEM) describing the direct and indirect effects of predictor variables on effect size of field.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1659478-g007.tif">
<alt-text content-type="machine-generated">Diagram illustrating the relationships between microbial count, precipitation, temperature, soil potassium, root length, potassium concentrations, leaf area, oxidase, and yield. Arrows indicate direction and strength of influence, with solid lines for significant relationships and dotted lines for insignificant ones. Values next to arrows represent correlation coefficients, and significance levels are marked with asterisks. The overall model fit is indicated by Chi-square, probability level, GFI, and AIC values.</alt-text>
</graphic>
</fig>
</sec>
</sec>
<sec id="s4" sec-type="discussion">
<label>4</label>
<title>Discussion</title>
<sec id="s4_1">
<label>4.1</label>
<title>Effects of KSM on different microbial genera and soil factors</title>
<p>Our results showed that KSM significantly increased crop yield, soil available K, and crop potassium uptake and growth, confirming their role in bioweathering potassium-containing minerals and mediating soil potassium transformation (<xref ref-type="bibr" rid="B4">Chen et&#xa0;al., 2022</xref>). Elevated root and shoot K concentrations further promote crop growth and yield, which aligns with Olaniyan who emphasized that enhanced potassium nutrition improves root vigor and photosynthetic efficiency (<xref ref-type="bibr" rid="B27">Olaniyan et&#xa0;al., 2022</xref>).</p>
<p>Crop yield and development varied with KSM genus, soil&#xa0;texture, and other contextual factors. Among tested microorganisms, <italic>Aspergillus</italic> showed the strongest overall effect on crop K content and yield, outperforming <italic>Bacillus</italic> and <italic>Pseudomonas</italic>. This superiority stems from its synergistic mechanisms: it secretes multiple extracellular organic acids (e.g., oxalic acid) to disrupt potassium-bearing mineral lattices and releases phosphatases/glucosidases (<xref ref-type="bibr" rid="B2">Ashrafi-Saiedlou et&#xa0;al., 2024</xref>), while producing phytohormones and iron transporters to optimize root physiological functions (<xref ref-type="bibr" rid="B18">Kumawat et&#xa0;al., 2024</xref>). Unlike <italic>Bacillus</italic> (which primarily relies on acidolysis), <italic>Aspergillus</italic> also improves rhizosphere microecology and soil aggregation (<xref ref-type="bibr" rid="B13">Imran et&#xa0;al., 2021</xref>), making it a key candidate for microbial preparation development and potash fertilizer efficiency improvement.</p>
<p>With respect to soil texture, clay loam showed the strongest yield response to KSM, which is attributed to the unique improvement effect of KSM on this soil type. Clay loam typically suffers from poor aeration that hinders root nutrient uptake (<xref ref-type="bibr" rid="B10">Hsiao et&#xa0;al., 2018</xref>), but KSM inoculation maintains stable available potassium levels during the crop reproductive period (<xref ref-type="bibr" rid="B34">Singh et&#xa0;al., 2022</xref>) and produces extracellular polysaccharides to promote soil aggregation (<xref ref-type="bibr" rid="B39">Wang et&#xa0;al., 2022</xref>). This structural improvement increases aeration and nutrient diffusion, enhancing root K uptake and root length (a trend consistent with the significant increase in root K concentration and length observed in clay loam). These findings indicate that KSM can targeted alleviate the limitations of clay loam soils, providing a solution for yield improvement in such agricultural systems.</p>
</sec>
<sec id="s4_2">
<label>4.2</label>
<title>Effects of KSM under different crop classifications, field management practices, and stress types</title>
<p>With respect to crop classification, our synthesis indicated that vegetable crops responded more positively to KSM than other crop types, which can be explained by their biological traits: most vegetables have short growth cycles, high potassium demand, and quality formation closely linked to potassium nutrition (<xref ref-type="bibr" rid="B44">Zoerb et&#xa0;al., 2014</xref>). Compared with cereals, vegetables such as sugar beet exhibit higher K mobilization efficiency per unit root length (<xref ref-type="bibr" rid="B7">El Dessougi et&#xa0;al., 2002</xref>; <xref ref-type="bibr" rid="B38">Wang et&#xa0;al., 2011</xref>), making them more sensitive to KSM-mediated potassium supplementation. This finding provides a basis for targeted KSM application&#x2014;prioritizing vegetable crops in agricultural production to maximize the return on microbial inoculation.</p>
<p>With respect to field management, greenhouse conditions significantly enhanced KSM efficacy relative to field trials, as the controlled temperature, humidity, and light optimize both crop growth and microbial survival (<xref ref-type="bibr" rid="B31">Shamshiri et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B8">Farvardin et&#xa0;al., 2024</xref>). This suggests that KSM application should be paired with appropriate environmental regulation to fully exploit its potential.</p>
<p>Under stress types, KSM consistently improved crop performance across drought and salinity gradients, which is achieved through dual mechanisms: supplementing potassium to maintain physiological functions (e.g., photosynthesis, enzyme activity) (<xref ref-type="bibr" rid="B25">Munsif et&#xa0;al., 2022</xref>) and enhancing stress defense systems (e.g., increasing antioxidant enzyme activity) (<xref ref-type="bibr" rid="B9">Feng et&#xa0;al., 2019</xref>). For salinity stress, KSM also helps maintain K<sup>+</sup>/Na<sup>+</sup> balance (<xref ref-type="bibr" rid="B28">Romero-Munar and Aroca, 2023</xref>), further confirming its role in improving crop stress resilience.</p>
</sec>
<sec id="s4_3">
<label>4.3</label>
<title>Key influences of the KSM on improving crop yields</title>
<p>This study demonstrated a significant linear correlation among soil available potassium, root/shoot potassium concentrations, and crop yield following KSM inoculation, consistent with previous findings (<xref ref-type="bibr" rid="B32">Shen et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B1">Ai et&#xa0;al., 2022</xref>). The increase in yield can be attributed to a well-defined physiological pathway initiated by KSM-mediated potassium solubilization.</p>
<p>KSM enhances the availability of soil potassium, leading to elevated potassium uptake by roots and subsequent translocation to shoots (<xref ref-type="bibr" rid="B3">Chen et&#xa0;al., 2021</xref>). The increased potassium levels stimulate root growth, including surface area, vigor, and biomass, which further expands nutrient absorption capacity and improves water retention (<xref ref-type="bibr" rid="B37">Sustr et&#xa0;al., 2019</xref>). Simultaneously, potassium promotes shoot development by facilitating cell expansion, leaf area enlargement, and photosynthetic efficiency (<xref ref-type="bibr" rid="B11">Hu et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B5">Ding et&#xa0;al., 2024</xref>).</p>
<p>Our SEM analysis revealed a multi-level regulatory network of KSM on yield, with soil available potassium and microbial count as core indirect drivers, and root length/leaf area as direct effectors. Soil available potassium promotes root tip cell division and elongation by regulating growth hormone transport (<xref ref-type="bibr" rid="B33">Sheng, 2005</xref>), while microbial count modulates root traits and nutrient availability (<xref ref-type="bibr" rid="B36">Sun et&#xa0;al., 2023</xref>), thereby establishing a coherent pathway from potassium activation to root growth and, ultimately, yield increase. Climate factors also play a role: MAP improves soil moisture to enhance root uptake (<xref ref-type="bibr" rid="B19">Li et&#xa0;al., 2023a</xref>), while MAT shows dual effects&#x2014;moderate temperatures promote metabolism, but extreme temperatures inhibit root K uptake (<xref ref-type="bibr" rid="B14">Jacobson et&#xa0;al., 1957</xref>), highlighting the need for temperature adaptation in KSM application.</p>
<p>These findings have important implications for sustainable agriculture: KSM can reduce reliance on mineral potash fertilizers&#xa0;by improving soil potassium use efficiency, particularly in clay loam and vegetable cropping systems. However, practical&#xa0;application must consider contextual limitations, for example, KSM efficacy may decline in highly saline soils due to microbial survival constraints. Future research should focus on optimizing inoculation strategies and developing climate-adapted KSM formulations to enhance applicability across diverse agricultural environments.</p>
</sec>
</sec>
<sec id="s5" sec-type="conclusions">
<label>5</label>
<title>Conclusions</title>
<p>This meta-analysis clarified the efficacy and regulatory mechanisms of KSM across diverse environments, filling the gap between local trials and global application guidelines. Key findings: (1) KSM inoculation significantly improved multiple agronomic parameters via a sequential process initiated by boosting soil available potassium, followed by enhanced plant potassium assimilation, and culminating in yield increase.; (2) Structural equation modeling identified microbial count and soil available K as core indirect drivers, with root length/leaf area as direct effectors; (3) <italic>Aspergillus</italic> was the most effective genus, with optimal responses in clay loam, vegetable crops, and greenhouse conditions. These findings provide a data-driven basis for reducing mineral potash fertilizer reliance&#x2014;practically, we recommend targeted inoculation of Aspergillus in clay loam vegetable fields, combined with greenhouse environmental regulation. In summary, KSM is a potent biological strategy for sustainable agriculture, and our work offers actionable pathways for its optimized application.</p>
</sec>
</body>
<back>
<sec id="s6" sec-type="data-availability">
<title>Data availability statement</title>
<p>The original contributions presented in the study are included in the article/<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Material</bold>
</xref>. Further inquiries can be directed to the corresponding author/s.</p>
</sec>
<sec id="s7" sec-type="author-contributions">
<title>Author contributions</title>
<p>XF: Writing &#x2013; original draft, Writing &#x2013; review &amp; editing. YZ:&#xa0;Writing &#x2013; original draft. YJ: Writing &#x2013; review &amp; editing. PD:&#xa0;Writing &#x2013; review &amp; editing. WW: Funding acquisition, Writing &#x2013; original draft. JL: Funding acquisition, Writing &#x2013; original draft. ZL: Funding acquisition, Writing &#x2013; original draft. RL: Writing &#x2013; original draft, Writing &#x2013; review &amp; editing. XL:&#xa0;Writing &#x2013; original draft.</p>
</sec>
<sec id="s8" sec-type="funding-information">
<title>Funding</title>
<p>The author(s) declare financial support was received for the&#xa0;research and/or publication of this article. This research was funded by the Key Research and Development Program Young Scientist Project (2023YFD1901900), the Natural Science Foundation of Shanxi province (202203021211139), the Inner Mongolia Ordos City Science and Technology Program, and Research and Promotion Project of Water Conservancy Science and Technology in Shanxi Province (2025GM15).</p>
</sec>
<sec id="s9" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="s10" sec-type="ai-statement">
<title>Generative AI statement</title>
<p>The author(s) declare that no Generative AI was used in the creation of this manuscript.</p>
<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&#xa0;you identify any issues, please contact us.</p>
</sec>
<sec id="s11" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<sec id="s12" sec-type="supplementary-material">
<title>Supplementary material</title>
<p>The Supplementary Material for this article can be found online at: <ext-link ext-link-type="uri" xlink:href="https://www.frontiersin.org/articles/10.3389/fpls.2025.1659478/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fpls.2025.1659478/full#supplementary-material</ext-link>
</p>
<supplementary-material xlink:href="Table1.xlsx" id="ST1" mimetype="application/vnd.openxmlformats-officedocument.spreadsheetml.sheet"/>
<supplementary-material xlink:href="DataSheet1.zip" id="SM1" mimetype="application/zip"/>
<supplementary-material xlink:href="DataSheet2.docx" id="SM2" mimetype="application/vnd.openxmlformats-officedocument.wordprocessingml.document"/>
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