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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Microbiol.</journal-id>
<journal-title>Frontiers in Microbiology</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Microbiol.</abbrev-journal-title>
<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.2025.1630650</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Microbiology</subject>
<subj-group>
<subject>Opinion</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Unlocking phosphorus resources: phosphate-solubilizing microorganisms as a green strategy for activating phosphorus in acidic red soils and promoting crop growth</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name><surname>Ma</surname> <given-names>Qingqing</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="http://loop.frontiersin.org/people/3068449/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/investigation/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Chen</surname> <given-names>Huayi</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/2781106/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Yang</surname> <given-names>Yuting</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="http://loop.frontiersin.org/people/3067992/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name><surname>Zhou</surname> <given-names>Bin</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>&#x0002A;</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/1918871/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/supervision/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
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</contrib-group>
<aff id="aff1"><sup>1</sup><institution>Tropical Crops Genetic Resources Institute, Chinese Academy of Tropical Agricultural Sciences, Haikou</institution>, <addr-line>Hainan</addr-line>, <country>China</country></aff>
<aff id="aff2"><sup>2</sup><institution>School of Tropical Agriculture and Forestry, Hainan University</institution>, <addr-line>Haikou</addr-line>, <country>China</country></aff>
<author-notes>
<fn fn-type="edited-by"><p>Edited by: Abubakar Dar, The Islamia University of Bahawalpur, Pakistan</p></fn>
<fn fn-type="edited-by"><p>Reviewed by: Zahir Ahmad Zahir, University of Agriculture, Faisalabad, Pakistan</p>
<p>Rubab Sarfraz, Gyeongsang National University, Republic of Korea</p></fn>
<corresp id="c001">&#x0002A;Correspondence: Bin Zhou <email>zhoubean&#x00040;vip.163.com</email></corresp>
</author-notes>
<pub-date pub-type="epub">
<day>09</day>
<month>07</month>
<year>2025</year>
</pub-date>
<pub-date pub-type="collection">
<year>2025</year>
</pub-date>
<volume>16</volume>
<elocation-id>1630650</elocation-id>
<history>
<date date-type="received">
<day>18</day>
<month>05</month>
<year>2025</year>
</date>
<date date-type="accepted">
<day>23</day>
<month>06</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x000A9; 2025 Ma, Chen, Yang and Zhou.</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>Ma, Chen, Yang and Zhou</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>
<kwd-group>
<kwd>microbial technology</kwd>
<kwd>red soil improvement</kwd>
<kwd>phosphorus inactivation</kwd>
<kwd>agricultural sustainability</kwd>
<kwd>tropical and subtropical agriculture</kwd>
</kwd-group>
<counts>
<fig-count count="1"/>
<table-count count="2"/>
<equation-count count="0"/>
<ref-count count="30"/>
<page-count count="5"/>
<word-count count="3316"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Microbiological Chemistry and Geomicrobiology</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec id="s1">
<title>1 Introduction</title>
<p>Tropical and subtropical agriculture is crucial for global food and economic crop production, and soil health directly impacts regional ecological security and food output. Red soils, widely distributed in these regions, are typical soil types formed by the long-term weathering of iron- and aluminum-rich oxides in hot and humid climates. Their red or brownish-red color results from the accumulation of iron and aluminum oxides. For instance, the red soil region in South China, covering 2.18 &#x000D7; 10<sup>6</sup> km<sup>2</sup> (about 36% of China&#x00027;s arable land), is key for economic and food crop production (Lang et al., <xref ref-type="bibr" rid="B15">2025</xref>; Huang and Zhao, <xref ref-type="bibr" rid="B10">2014</xref>). However, due to their strong acidity, low base saturation, and high phosphorus (P) fixation capacity, red soils are among the most vulnerable soil types in agricultural production (Long et al., <xref ref-type="bibr" rid="B19">2024</xref>).</p>
<p>P is essential for plant growth and development, driving cell division, differentiation, and metabolic processes (Ashley et al., <xref ref-type="bibr" rid="B2">2011</xref>). While weathering of red soil parent material releases a significant amount of P, its bioavailability is constrained by the unique formation process of soil and mineral characteristics, resulting in a low flux biogeochemical cycle of P in red soil ecosystems (Ma et al., <xref ref-type="bibr" rid="B21">2016</xref>). Under acidic conditions, secondary clay minerals and amorphous iron and aluminum oxides expose numerous hydroxyl sites, which adsorb phosphate ions specifically, forming stable inner-sphere complexes. Consequently, P is fixed in soil solids as amorphous or crystalline iron- and aluminum-phosphate (He et al., <xref ref-type="bibr" rid="B7">2025</xref>). This chemical fixation is largely irreversible, creating a surplus of fixed P and significantly limiting the flux of available P in the soil-plant system (Huang et al., <xref ref-type="bibr" rid="B11">2021</xref>). Moreover, the low organic matter content in red soils weakens the P coordination-dissolution balance. Soil aggregate destruction caused by intensive farming further exacerbates P leaching into deep soil layers (Huang et al., <xref ref-type="bibr" rid="B11">2021</xref>). These factors, which cause P deactivation, severely restrict agricultural production in red soils. Critically, sustainable alternatives remain underexplored, particularly regarding microbiome-driven P activation. This review uniquely bridges microbial ecology with agronomic application, highlighting how phosphate-solubilizing microorganisms (PSMs) concurrently resolve P limitation, aluminum toxicity, and soil structural decline-gaps unaddressed by chemical approaches.</p>
</sec>
<sec id="s2">
<title>2 PSMs have unique potential in red soil improvement</title>
<sec>
<title>2.1 Microbial technologies are widely used in red soil improvement</title>
<p>In the global context of sustainable agriculture, microbial technology, known for being eco-friendly and functionally diverse, has become a key research focus in red soil improvement. Traditional chemical methods, such as lime application and chemical phosphate fertilizer addition, can temporarily adjust soil acidity or supplement P (<xref ref-type="table" rid="T1">Table 1</xref>). However, their long-term use may lead to problems like secondary salinization, soil structure degradation, and increased ecological risks (Ji et al., <xref ref-type="bibr" rid="B12">2024</xref>). In contrast, microbial technology can regulate soil biogeochemical cycles, thereby achieving multiple goals of nutrient activation, toxicity mitigation, and ecological restoration. Microbial metabolism directly acts on soil minerals and organic matter, releasing fixed nutrients and enhancing the soil microenvironment via metabolic products (Khan et al., <xref ref-type="bibr" rid="B13">2024</xref>). Additionally, microbial growth promotes soil aggregate formation, thereby improving soil structure and boosting water and nutrient retention capacities (Long et al., <xref ref-type="bibr" rid="B20">2025</xref>).</p>
<table-wrap position="float" id="T1">
<label>Table 1</label>
<caption><p>Methods of red soil improvement and their advantages and drawbacks.</p></caption>
<table frame="box" rules="all">
<thead>
<tr style="background-color:#919498;color:#ffffff">
<th valign="top" align="left"><bold>Types</bold></th>
<th valign="top" align="left"><bold>Technologies</bold></th>
<th valign="top" align="left"><bold>Advantages</bold></th>
<th valign="top" align="left"><bold>Drawbacks</bold></th>
<th valign="top" align="left"><bold>References</bold></th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left" rowspan="4">Chemical</td>
<td valign="top" align="left">Liming</td>
<td valign="top" align="left">&#x02022; Quickly neutralizes soil acidity<break/> &#x02022; Reduces aluminum toxicity</td>
<td valign="top" align="left">&#x02022; Excessive use can lead to soil compaction</td>
<td valign="top" align="left">Xu et al. (<xref ref-type="bibr" rid="B29">2023</xref>)</td>
</tr>
 <tr>
<td valign="top" align="left">Oyster shell Powder</td>
<td valign="top" align="left">&#x02022; Neutralizes soil acidity<break/> &#x02022; Supplies calcium and magnesium</td>
<td valign="top" align="left">&#x02022; Slow improvement rate<break/> &#x02022; Influenced by soil conditions</td>
<td valign="top" align="left">Li et al. (<xref ref-type="bibr" rid="B16">2025</xref>)</td>
</tr>
 <tr>
<td valign="top" align="left">Phosphogypsum</td>
<td valign="top" align="left">&#x02022; Neutralizes acidity<break/> &#x02022; Provides calcium and sulfur</td>
<td valign="top" align="left">&#x02022; Contains heavy metals and harmful substances</td>
<td valign="top" align="left">Sun et al. (<xref ref-type="bibr" rid="B27">2025</xref>)</td>
</tr>
 <tr>
<td valign="top" align="left">Biochar</td>
<td valign="top" align="left">&#x02022; Neutralizes acidity<break/> &#x02022; Improves soil fertility</td>
<td valign="top" align="left">&#x02022; Affects soil infiltration<break/> &#x02022; Uncertain long-term effects</td>
<td valign="top" align="left">He et al. (<xref ref-type="bibr" rid="B7">2025</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Physical</td>
<td valign="top" align="left">Deep tillage</td>
<td valign="top" align="left">&#x02022; Enhances soil aeration and permeability<break/> &#x02022; Promotes microbial activity</td>
<td valign="top" align="left">&#x02022; Mechanical compaction may cause local soil compaction</td>
<td valign="top" align="left">Chen et al. (<xref ref-type="bibr" rid="B3">2020</xref>)</td>
</tr>
<tr>
<td valign="top" align="left" rowspan="2">Biological</td>
<td valign="top" align="left">Microbial inoculant</td>
<td valign="top" align="left">&#x02022; Increases beneficial microorganisms<break/> &#x02022; Promotes nutrient transformation</td>
<td valign="top" align="left">&#x02022; Microbial growth is affected by environmental conditions</td>
<td valign="top" align="left">Shi et al. (<xref ref-type="bibr" rid="B26">2024</xref>)</td>
</tr>
 <tr>
<td valign="top" align="left">Plant-based remediation</td>
<td valign="top" align="left">&#x02022; Lowers heavy metal toxicity<break/> &#x02022; Improves soil ecosystem</td>
<td valign="top" align="left">&#x02022; Long restoration period<break/> &#x02022; Plant species selection is crucial</td>
<td valign="top" align="left">Li et al. (<xref ref-type="bibr" rid="B17">2024</xref>)</td>
</tr>
<tr>
<td valign="top" align="left" rowspan="2">Agronomic Practices</td>
<td valign="top" align="left">Organic manure</td>
<td valign="top" align="left">&#x02022; Raises organic matter content<break/> &#x02022; Improves soil structure</td>
<td valign="top" align="left">&#x02022; Slow nutrient release<break/> &#x02022; Fails to meet immediate crop needs</td>
<td valign="top" align="left">Qiu et al. (<xref ref-type="bibr" rid="B24">2025</xref>)</td>
</tr>
 <tr>
<td valign="top" align="left">Crop rotation</td>
<td valign="top" align="left">&#x02022; Improves soil physical and chemical properties<break/> &#x02022; Reduces pests and diseases</td>
<td valign="top" align="left">&#x02022; Rotation mode selection is complex</td>
<td valign="top" align="left">Mao et al. (<xref ref-type="bibr" rid="B22">2025</xref>)</td>
</tr></tbody>
</table>
</table-wrap>
</sec>
<sec>
<title>2.2 PSMs as potential candidates for red soil amelioration</title>
<p>PSMs are beneficial functional microorganisms that convert insoluble environmental phosphorus into plant-available forms. They were first identified in farmland and environmental studies (Vassilev et al., <xref ref-type="bibr" rid="B28">2006</xref>). PSMs secrete acids and extracellular enzymes to dissolve insoluble inorganic and organic phosphates. Additionally, they release intracellular P through cell lysis, thereby enhancing soil P availability and improving soil microbial community balance (Hu and Chen, <xref ref-type="bibr" rid="B9">2023</xref>). These characteristics provide PSMs with a unique advantage in red soil improvement. Notably, their ecological adaptability enables them to remain active under the extremely acidic conditions of red soils. For instance, studies have shown that Pseudomonas duriflava, Enterobacter quasimori, and Acinetobacter sp. exhibit robust phosphorus-solubilizing abilities at pH 4.5&#x02013;6.2, indicating their genetic potential to adapt to red soil acidification (Kumar et al., <xref ref-type="bibr" rid="B14">2025</xref>; Hidayat et al., <xref ref-type="bibr" rid="B8">2024</xref>; Liu et al., <xref ref-type="bibr" rid="B18">2014</xref>). Importantly, PSMs can maintain soil P availability over time, providing sustained nutrient support for crop growth, reducing reliance on external phosphate fertilizers, and thus aligning with the goals of red soil improvement and sustainable agriculture (Rawat et al., <xref ref-type="bibr" rid="B25">2021</xref>).</p>
</sec>
</sec>
<sec id="s3">
<title>3 PSMs regulate community structure to enhance red soil fertility via P activation</title>
<p>PSMs improve red soil fertility not only through their inherent functions but also by regulating the soil microbial community to form a synergistic ecological network (<xref ref-type="fig" rid="F1">Figure 1</xref>). The decline in microbial diversity and functional imbalance caused by red soil acidification are key factors limiting nutrient cycling. Introduced PSMs can reshape the rhizosphere microbiome while modifying the local microenvironment. For example, they promote the colonization of beneficial bacteria, such as nitrogen-fixing bacteria, and boost the relative abundance of function microbes related to P cycling (Rawat et al., <xref ref-type="bibr" rid="B25">2021</xref>). This community optimization builds a metabolic module centered on P activation, thereby strengthening the transformation efficiency of soil nutrients. Specifically, organic acids, inorganic acids, and enzymes secreted by PSMs can solubilize occluded P by protonating Fe-P and Al-P surfaces, thereby increasing available soil P and alleviating aluminum toxicity (Ahmad et al., <xref ref-type="bibr" rid="B1">2023</xref>). Moreover, PSM metabolic products (e.g., exopolysaccharides, siderophores) stabilize soil aggregates, thereby improving pore structure and water-holding capacity and creating a favorable habitat for microbial activity (Khan et al., <xref ref-type="bibr" rid="B13">2024</xref>).</p>
<fig id="F1" position="float">
<label>Figure 1</label>
<caption><p>Schematic diagram of phosphate-solubilizing microorganisms improving red soil and crop growth.</p></caption>
<alt-text>Diagram contrasting phosphorus deactivation and solubilization in soil. The left side shows aluminum toxicity causing cellular destruction and cytokenesis impairment in red soil, with insoluble phosphorus bound with iron and aluminum. The right side illustrates phosphate solubilizing microorganisms (PSMs) enhancing root growth and rhizosphere community, reducing ethylene, and improving soil structure through enzyme and acid release. Labels indicate processes like phosphorus dissolution and soil microbial regulation.</alt-text>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmicb-16-1630650-g0001.tif"/>
</fig>
</sec>
<sec id="s4">
<title>4 PSMs boost crop yield and stress resistance in red soils through multifaceted promotion</title>
<p>The effects of PSMs on red soil improvement and crop promotion are presented in <xref ref-type="table" rid="T2">Table 2</xref>. The growth-promoting effects of PSMs on red soil crops extend beyond mere nutrient supply. By multidimensionally regulating plant physiology, PSMs enhance crops&#x00027; environmental adaptability, ensuring stable and increased yields under stress. Under red soil&#x00027;s strong acidity and aluminum toxicity, PSMs secrete plant hormones such as indoleacetic acid and gibberellins. These hormones stimulate root meristem activity, promoting lateral root and root hair development, thereby expanding nutrient absorption (Masrahi et al., <xref ref-type="bibr" rid="B23">2023</xref>; Feng et al., <xref ref-type="bibr" rid="B5">2024</xref>). Meanwhile, the ACC deaminase produced by PSMs degrades the ethylene precursor generated under plant stress, thereby alleviating root growth inhibition (Kumar et al., <xref ref-type="bibr" rid="B14">2025</xref>). When confronting oxidative stress, PSMs activate the antioxidant enzyme system in crops (e.g., superoxide dismutase, peroxidase), scavenging excess reactive oxygen species and maintaining cell membrane stability (Rawat et al., <xref ref-type="bibr" rid="B25">2021</xref>). Furthermore, the mutualistic relationship between PSMs and crop roots can induce systemic resistance in plants, thereby strengthening defense against soil-borne pathogens. For instance, combined amino acid and PSM applications have reduced pathogenic fungal relative abundance by 5.2%, thereby lowering disease risk (Shi et al., <xref ref-type="bibr" rid="B26">2024</xref>). This mutualistic relationship involves complex signaling pathways and metabolic interactions. PSMs produce specific molecules that trigger plant defense responses, such as the production of pathogenesis-related proteins and secondary metabolites (Guo et al., <xref ref-type="bibr" rid="B6">2024</xref>). In field practices, PSM applications consistently improve crop growth, foliar nitrogen and phosphorus concentrations, grain quality, and yields (Masrahi et al., <xref ref-type="bibr" rid="B23">2023</xref>). This comprehensive regulation, ranging from soil improvement to enhancing crop stress resistance, highlights PSMs&#x00027; central value in red soil agro-ecosystems.</p>
<table-wrap position="float" id="T2">
<label>Table 2</label>
<caption><p>Phosphorus solubilizing capacity of PSMs and its effect on red soil improvement and crop growth.</p></caption>
<table frame="box" rules="all">
<thead>
<tr style="background-color:#919498;color:#ffffff">
<th valign="top" align="left"><bold>PSM strains</bold></th>
<th valign="top" align="left"><bold>Genus</bold></th>
<th valign="top" align="center"><bold>P solubility (mg/L)</bold></th>
<th valign="top" align="center"><bold>Soil available P improvement</bold></th>
<th valign="top" align="left"><bold>Crop types</bold></th>
<th valign="top" align="left"><bold>Crop growth</bold></th>
<th valign="top" align="left"><bold>References</bold></th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">ASL12</td>
<td valign="top" align="left"><italic>Acinetobacter</italic> sp.</td>
<td valign="top" align="center">717</td>
<td valign="top" align="center">51.27%</td>
<td valign="top" align="left"><italic>Areca catechu</italic></td>
<td valign="top" align="left">82.84% increase in plant height</td>
<td valign="top" align="left">Liu et al. (<xref ref-type="bibr" rid="B18">2014</xref>)</td>
</tr>
 <tr>
<td valign="top" align="left">ASG33</td>
<td valign="top" align="left"><italic>Shigella</italic> sp.</td>
<td valign="top" align="center">530</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">/</td>
<td valign="top" align="left">/</td>
<td/>
</tr>
 <tr>
<td valign="top" align="left">ASG34</td>
<td valign="top" align="left"><italic>Escherichia</italic> sp.</td>
<td valign="top" align="center">499</td>
<td valign="top" align="center">34.32%</td>
<td valign="top" align="left"><italic>Areca catechu</italic></td>
<td valign="top" align="left">74.90% increase in plant height</td>
<td/>
</tr>
 <tr>
<td valign="top" align="left">ADH302</td>
<td valign="top" align="left"><italic>Enterobacter</italic> sp.</td>
<td valign="top" align="center">426</td>
<td valign="top" align="center">24.15%</td>
<td valign="top" align="left"><italic>Areca catechu</italic></td>
<td valign="top" align="left">71.55% increase in plant height</td>
<td/>
</tr>
 <tr>
<td valign="top" align="left">ATZ304</td>
<td valign="top" align="left"><italic>Paenibacillus</italic> sp.</td>
<td valign="top" align="center">266</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">/</td>
<td valign="top" align="left">/</td>
<td/>
</tr>
 <tr>
<td valign="top" align="left">ASG64</td>
<td valign="top" align="left"><italic>Paenibacillus</italic> sp.</td>
<td valign="top" align="center">165</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">/</td>
<td valign="top" align="left">/</td>
<td/>
</tr>
 <tr>
<td valign="top" align="left">ADH306</td>
<td valign="top" align="left"><italic>Bacillus</italic> sp.</td>
<td valign="top" align="center">137</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">/</td>
<td valign="top" align="left">/</td>
<td/>
</tr>
 <tr>
<td valign="top" align="left">ASG16</td>
<td valign="top" align="left"><italic>Kurthia</italic> sp.</td>
<td valign="top" align="center">439</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">/</td>
<td valign="top" align="left">/</td>
<td/>
</tr>
 <tr>
<td valign="top" align="left">ASG41</td>
<td valign="top" align="left"><italic>Paenibacillus</italic> sp.</td>
<td valign="top" align="center">300</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">/</td>
<td valign="top" align="left">/</td>
<td/>
</tr>
 <tr>
<td valign="top" align="left">AX7</td>
<td valign="top" align="left"><italic>Rhizobium</italic> sp.</td>
<td valign="top" align="center">275</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">/</td>
<td valign="top" align="left">/</td>
<td/>
</tr>
<tr>
<td valign="top" align="left">NBW</td>
<td valign="top" align="left"><italic>Bacillus subtilis</italic></td>
<td valign="top" align="center">/</td>
<td valign="top" align="center">81.0%</td>
<td valign="top" align="left">Pepper</td>
<td valign="top" align="left">66.50% increase in yield</td>
<td valign="top" align="left">Duan et al. (<xref ref-type="bibr" rid="B4">2024</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">PSB 5</td>
<td valign="top" align="left">/</td>
<td valign="top" align="center">44.4</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">/</td>
<td valign="top" align="left">/</td>
<td valign="top" align="left">Kumar et al. (<xref ref-type="bibr" rid="B14">2025</xref>)</td>
</tr>
 <tr>
<td valign="top" align="left">PSB 8</td>
<td valign="top" align="left">/</td>
<td valign="top" align="center">48.0</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">/</td>
<td valign="top" align="left">/</td>
<td/>
</tr>
 <tr>
<td valign="top" align="left">PSB 9</td>
<td valign="top" align="left">/</td>
<td valign="top" align="center">44.7</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">/</td>
<td valign="top" align="left">/</td>
<td/>
</tr>
 <tr>
<td valign="top" align="left">PSB 10</td>
<td valign="top" align="left">/</td>
<td valign="top" align="center">53.4</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">/</td>
<td valign="top" align="left">/</td>
<td/>
</tr>
<tr>
<td valign="top" align="left">13-2</td>
<td valign="top" align="left"><italic>Bacillus amyloliquefaciens</italic></td>
<td valign="top" align="center">/</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">Tomato</td>
<td valign="top" align="left">21.38% increase in plant height</td>
<td valign="top" align="left">Guo et al. (<xref ref-type="bibr" rid="B6">2024</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">RW37</td>
<td valign="top" align="left"><italic>Enterobacter soli</italic></td>
<td valign="top" align="center">498</td>
<td valign="top" align="center">/</td>
<td valign="top" align="left">Moso bamboo</td>
<td valign="top" align="left">56.49% increase in plant height</td>
<td valign="top" align="left">Zhang et al. (<xref ref-type="bibr" rid="B30">2024</xref>)</td>
</tr></tbody>
</table>
</table-wrap>
</sec>
<sec id="s5">
<title>5 Future research directions and strategies</title>
<p>PSMs activate soil P reserves, alleviate aluminum toxicity, and secrete growth promoters, representing a cost-effective and sustainable green strategy for red soil improvement. However, given some limitations of PSMs, future research can be further deepened in the following aspects: Firstly, targeting the extreme acidity of red soil, screening and engineering functional strains with efficient P solubilization and acid tolerance is crucial. Decoding the acid-resistant molecular mechanisms of PSMs via genomics and metabolomics, combined with gene editing to enhance their phosphorus-solubilizing capacity or aluminum detoxification pathways, can boost strain adaptability. Secondly, developing integrated restoration systems that combine multiple technologies is essential to overcome the limitations of single microbial technologies. For example, combining PSMs with biochar, phosphate minerals, and harmless phosphogypsum can provide microbial habitats and synergistically regulate soil pH. Additionally, it is necessary to comprehensively evaluate the long-term impacts of PSM application on the structure and functions of soil microbial communities. This can prevent potential imbalances or functional redundancies in indigenous microbial populations that may be caused by the introduction of exogenous microorganisms.</p>
</sec>
</body>
<back>
<sec sec-type="author-contributions" id="s6">
<title>Author contributions</title>
<p>QM: Investigation, Writing &#x02013; original draft. HC: Writing &#x02013; review &#x00026; editing. YY: Writing &#x02013; original draft. BZ: Supervision, Writing &#x02013; review &#x00026; editing, Funding acquisition.</p>
</sec>
<sec sec-type="funding-information" id="s7">
<title>Funding</title>
<p>The author(s) declare that financial support was received for the research and/or publication of this article. This study was supported by the National Key Research and Development Program of China (2023YFD1901303) and Central Public Interest Scientific Institution Basal Research Fund (1630032023010, Tropical Crops Genetic Resources Institute, CATAS).</p>
</sec>
<sec sec-type="COI-statement" id="conf1">
<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 sec-type="ai-statement" id="s8">
<title>Generative AI statement</title>
<p>The author(s) declare that no Gen AI was used in the creation of this manuscript.</p>
</sec>
<sec sec-type="disclaimer" id="s9">
<title>Publisher&#x00027;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>
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