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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.2025.1466452</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Microbiology</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Impact of root exudates on soil reconstruction and bacterial community resumption in open-pit coal mines</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name><surname>Yang</surname> <given-names>Zhuo</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
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<contrib contrib-type="author" corresp="yes">
<name><surname>Niu</surname> <given-names>Jianzhi</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
<xref ref-type="aff" rid="aff4"><sup>4</sup></xref>
<xref ref-type="aff" rid="aff5"><sup>5</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>&#x002A;</sup></xref>
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<contrib contrib-type="author">
<name><surname>Wu</surname> <given-names>Tong</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
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<contrib contrib-type="author">
<name><surname>Li</surname> <given-names>Jiaqi</given-names></name>
<xref ref-type="aff" rid="aff6"><sup>6</sup></xref>
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<contrib contrib-type="author">
<name><surname>Zhang</surname> <given-names>Linus</given-names></name>
<xref ref-type="aff" rid="aff7"><sup>7</sup></xref>
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<contrib contrib-type="author">
<name><surname>Chen</surname> <given-names>Xiongwen</given-names></name>
<xref ref-type="aff" rid="aff8"><sup>8</sup></xref>
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<contrib contrib-type="author">
<name><surname>Berndtsson</surname> <given-names>Ronny</given-names></name>
<xref ref-type="aff" rid="aff7"><sup>7</sup></xref>
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<aff id="aff1"><sup>1</sup><institution>School of Soil and Water Conservation, Beijing Forestry University</institution>, <addr-line>Beijing</addr-line>, <country>China</country></aff>
<aff id="aff2"><sup>2</sup><institution>China Coal Technology &#x0026; Engineering Group Shenyang Engineering Company</institution>, <addr-line>Shenyang</addr-line>, <country>China</country></aff>
<aff id="aff3"><sup>3</sup><institution>State Key Laboratory of Efficient Production of Forest Resources</institution>, <addr-line>Beijing</addr-line>, <country>China</country></aff>
<aff id="aff4"><sup>4</sup><institution>Key Laboratory of State Forestry Administration on Soil and Water Conservation and Desertification Combating</institution>, <addr-line>Beijing</addr-line>, <country>China</country></aff>
<aff id="aff5"><sup>5</sup><institution>Engineering Research Center of Forestry Ecological Engineering of Ministry of Education, Beijing Forestry University</institution>, <addr-line>Beijing</addr-line>, <country>China</country></aff>
<aff id="aff6"><sup>6</sup><institution>College of Land and Environment, Shenyang Agricultural University</institution>, <addr-line>Shenyang</addr-line>, <country>China</country></aff>
<aff id="aff7"><sup>7</sup><institution>Division of Water Resources Engineering and Centre for Advanced Middle Eastern Studies, Lund University</institution>, <addr-line>Lund</addr-line>, <country>Sweden</country></aff>
<aff id="aff8"><sup>8</sup><institution>Department of Biological and Environmental Sciences, Alabama A&#x0026;M University</institution>, <addr-line>Huntsville, AL</addr-line>, <country>United States</country></aff>
<author-notes>
<fn fn-type="edited-by" id="fn0002">
<p>Edited by: Samina Mehnaz, Forman Christian College, Pakistan</p>
</fn>
<fn fn-type="edited-by" id="fn0003">
<p>Reviewed by: Chunqiao Xiao, Wuhan Institute of Technology, China</p>
<p>Shiliang Liu, Beijing Normal University, China</p>
</fn>
<corresp id="c001">&#x002A;Correspondence: Jianzhi Niu, <email>nexk@bjfu.edu.cn</email></corresp>
</author-notes>
<pub-date pub-type="epub">
<day>30</day>
<month>07</month>
<year>2025</year>
</pub-date>
<pub-date pub-type="collection">
<year>2025</year>
</pub-date>
<volume>16</volume>
<elocation-id>1466452</elocation-id>
<history>
<date date-type="received">
<day>18</day>
<month>07</month>
<year>2024</year>
</date>
<date date-type="accepted">
<day>02</day>
<month>07</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x00A9; 2025 Yang, Niu, Wu, Li, Zhang, Chen and Berndtsson.</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>Yang, Niu, Wu, Li, Zhang, Chen and Berndtsson</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>Open-pit coal mine reconstructed ecosystems are ecologically fragile. Retained early stage topsoil is usually not enough to maintain plant growth. For this purpose, we used root exudates to fertilize the reconstructed soil and improve the functioning of the soil microorganism ecology. The roots&#x2019; exudates increased the concentration of organic matter and total nitrogen by 16&#x2013;39%. Within a certain concentration range, the higher the concentration of root exudate, the higher the soil fertility. When the concentration of root exudate was 85%, the bacterial abundance decreased. The soil inorganic nitrogen N-NH<sub>4</sub><sup>+</sup> and N-NO<sub>3</sub><sup>&#x2212;</sup> increased significantly by 11&#x2013;21%. This significantly improved root growth and plant biomass for the reconstructed soil. The dominating bacterial community was driven by both root exudate components and plant root growth. Especially, the abundance of soil bacteria Actinobacteriota, Proteobacteria, and Chloroflexi was significantly promoted. Consequently, root exudates can be used to efficiently increase the soil fertility and improve the function and vegetation restoration in the soil reconstruction of mines.</p>
</abstract>
<kwd-group>
<kwd>root exudates</kwd>
<kwd>open-pit coal mines</kwd>
<kwd>soil reconstruction</kwd>
<kwd>bacteria community</kwd>
<kwd>diversity and abundance</kwd>
</kwd-group>
<counts>
<fig-count count="9"/>
<table-count count="2"/>
<equation-count count="0"/>
<ref-count count="81"/>
<page-count count="13"/>
<word-count count="8832"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Terrestrial Microbiology</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec sec-type="intro" id="sec1">
<label>1</label>
<title>Introduction</title>
<p>Open-pit mining is globally the main coal mining method. Many countries have realized the importance of land reclamation and have successively promulgated laws and regulations with respect to land reclamation of abandoned mining sites. Soil reconstruction is the core of land reclamation, and the quality of soil reconstruction directly determines the land reclamation status (<xref ref-type="bibr" rid="ref60">Wang et al., 2021</xref>). The original topsoil is the best choice to reconstruct the soil. However, the poor soil development in mining areas due to mining activities causes serious problems of surface soil scarcity (<xref ref-type="bibr" rid="ref48">Robson et al., 2018</xref>). When topsoil is scarce, suitable soil substitute materials are often screened from the geotechnical layer above the coal seam (<xref ref-type="bibr" rid="ref43">Nicolini and Topp, 2005</xref>; <xref ref-type="bibr" rid="ref63">Wilson-Kokes et al., 2013</xref>). Open-pit coal mines in China are mostly located in arid and semi-arid areas. Numerous domestic and foreign research results have shown that the native soil in open-pit mines is deficient in nutrients, and the soil fertility and plant diversity are poor, which is insufficient to realize the functions of soil nutrient cycling and microbial regulation (<xref ref-type="bibr" rid="ref61">Wang et al., 2022</xref>; <xref ref-type="bibr" rid="ref15">Feng, 2019</xref>; <xref ref-type="bibr" rid="ref9003">Hamid et al., 2022</xref>). The types of green vegetation available for ecological restoration are limited (<xref ref-type="bibr" rid="ref64">Wu et al., 2023</xref>; <xref ref-type="bibr" rid="ref70">Zhao et al., 2023</xref>). Thus, it is necessary to find an amendment for surface reconstruction soil that can balance the requirements of soil microorganism reproduction and the growth needs of locally adapted plants.</p>
<p>Root exudates are the general term for organic compounds released from the roots to the growth medium during the growth and development of plants, which can generally account for more than 10% of the photosynthetic carbon sequestration of plants (<xref ref-type="bibr" rid="ref17">Haichar et al., 2016</xref>). Root exudates include low molecular weight organic substances, high molecular weight mucilage substances, root cell abscission, and its decomposition products, as well as gases, protons, and nutrient ions (<xref ref-type="bibr" rid="ref24">Jiang et al., 2022</xref>). They are key factors in regulating the micro-ecological functions of the rhizosphere and also the main medium for communication between plants and soil (<xref ref-type="bibr" rid="ref7">Chen et al., 2014</xref>; <xref ref-type="bibr" rid="ref45">Pausch and Kuzyakov, 2018</xref>). They are of great significance for the elemental cycling of soil in the reclamation of mining areas, the absorption of plant nutrients, and the improvement of microbial functions (<xref ref-type="bibr" rid="ref55">Ulbrich et al., 2022</xref>; <xref ref-type="bibr" rid="ref11">Damin et al., 2008</xref>). A large number of previous empirical studies have shown that the growth of plants under the influence of root exudates is relatively good (<xref ref-type="bibr" rid="ref9001">Carla et al., 2020</xref>; <xref ref-type="bibr" rid="ref57">Viviane et al., 2021</xref>). To adapt to the soil environment, plants will adjust the microbial community&#x2019;s absorption and transformation of nutrient elements by changing the secretion of their root systems (<xref ref-type="bibr" rid="ref46">Qu et al., 2020</xref>; <xref ref-type="bibr" rid="ref51">Sun et al., 2021</xref>; <xref ref-type="bibr" rid="ref58">Wang et al., 2023</xref>; <xref ref-type="bibr" rid="ref47">Rahman et al., 2021</xref>). Plant root secretion is influenced by biological factors (plant species genotype) and abiotic factors (soil properties, climatic conditions) (<xref ref-type="bibr" rid="ref23">Huang et al., 2019</xref>; <xref ref-type="bibr" rid="ref22">Hu et al., 2021</xref>). The components of root exudates secreted by different kinds of plants grown under specific soil conditions are variable (<xref ref-type="bibr" rid="ref1">Alberto et al., 2019</xref>; <xref ref-type="bibr" rid="ref52">Sun et al., 2018</xref>). In the rhizosphere soil, bacteria have an absolute preponderant number of rhizosphere microorganisms, which plays an important role in maintaining plant growth and reducing the influence of environmental stress on plants (<xref ref-type="bibr" rid="ref26">Jones et al., 2017</xref>; <xref ref-type="bibr" rid="ref40">Meihong et al., 2019</xref>). Plant roots affect the population structure and diversity of rhizosphere bacteria by regulating the type and quantity of root exudates (<xref ref-type="bibr" rid="ref9">Cordovez et al., 2024</xref>). Rhizosphere bacteria regulate the secretion of plant roots by regulating soil nutrients, thus affecting the growth and development of plants and nutrient absorption (<xref ref-type="bibr" rid="ref37">Lyu et al., 2024</xref>). The nutrient content, transformation, and utilization efficiency of thin reconstructed soil in mining areas are often low, and the biological adaptability of the reconstructed soil is poor (<xref ref-type="bibr" rid="ref27">Ju et al., 2021</xref>; <xref ref-type="bibr" rid="ref12">Demyan and Smeck, 2022</xref>). According to the above, a way to overcome some of these problems may be to use root exudates to recover soil fertility and microbial diversity. It has been shown that low soil microbial biomass can restrict the recovery of soil functioning during the restoration of post-mining lands in semi-arid regions. Thus, especially for mining areas in areas with naturally nutrient-poor soils in arid and semiarid regions, this could be helpful in plant nutrients&#x2019; absorption, transformation, and utilization, as an organic modifier and assist microorganism breeding and ecological function recovering in the reconstruction soil land (<xref ref-type="bibr" rid="ref36">Lu et al., 2018</xref>; <xref ref-type="bibr" rid="ref9004">Li et al., 2019</xref>). The research team selected the typical open-pit mine in China - Zahanoer open-pit mine, Inner Mongolia as the research object. Based on the investigation results, sweet clover (<italic>Melilotus officinalis</italic> L. Pall) was selected as the root exudate extraction body because it is a common reclamation species in mines and has high economic efficiency. Based on the research findings regarding the growth characteristics of common reclamation plants in open-pit mines and in accordance with experimental requirements, it is determined that <italic>Medicago sativa</italic> (<italic>Medicago Sativa Linn</italic>) exhibits significant salt-alkali tolerance and drought resistance. Its genetic mechanism provides a biological foundation for soil improvement in open-pit mines. <italic>Medicago sativa</italic> demonstrates strong adaptability to the reclaimed areas of open-pit mines. As the &#x201C;king of forage grass, &#x201C;its cultivation techniques are well-established, and extensive large-scale application experience has been accumulated in arid and semi-arid regions such as Inner Mongolia. Moreover, the molecular mechanisms and field monitoring technologies related to <italic>Medicago sativa</italic> have developed into a relatively comprehensive research system, facilitating experimental design and data collection. Consequently, this study ultimately selects <italic>Medicago sativa</italic>, a pioneer herbaceous reclamation plant, as the experimental subject to investigate the functions of root secretion improvement and microbial regulation. The results can be used to provide basic technical support for ideas and inspiration in the ecological restoration of opencast coal mine areas.</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>Preparation of root exudates</title>
<p>The overburden of the seam in Zhalute County open-pit coal mine, Inner Mongolia, China (120.91507&#x00B0;E, 44.55592&#x00B0;N), was used to reconstruct topsoil and plant herbage. The reconstruction soil was made up of carbonaceous mudstone, mudstone, coal seam floor earth, subsoil, and sandy loam in a weight ratio of 2:1:2:2:2, respectively. The reconstruction soil contained total carbon (TC) of 148&#x202F;g&#x202F;kg<sup>&#x2212;1</sup>, total nitrogen (TN) of 1.5&#x202F;g&#x202F;kg<sup>&#x2212;1</sup>, bulk density of 1.32&#x202F;g&#x00B7;cm<sup>&#x2212;3</sup>, pH of 7.36, and a particle size of 0.5&#x2013;3&#x202F;mm. No farmland soil was used in the process.</p>
<p>To obtain root exudates, we cultivated sweet clover (<italic>Melilotus officinalis</italic> L. Pall.) in a sterile environment. The seeds undergo disinfection via a stepwise disinfection method. Specifically, they were first soaked in 75% ethanol for 2&#x202F;min, followed by immersion in a 3% H<sub>2</sub>O<sub>2</sub> solution for 15&#x202F;min, and subsequently rinsed five times with sterile water to thoroughly inactivate any resistant microorganisms present on or within the seeds. This process effectively prevents contamination during subsequent cultivation. Thereafter, the seeds were planted in cups containing a sponge that had been pre-treated by sterilization at 121&#x00B0;Cunder high pressure for 30&#x202F;min and dried in a 60&#x00B0;C oven for 24&#x202F;h to eliminate potential microbial growth caused by condensation. The <italic>Hoagland Nutrient Solution</italic> provided nutrition for plant growth (<xref ref-type="table" rid="tab1">Table 1</xref>). When preparing the nutrient solution, it was filtered and sterilized using a 0.22&#x202F;&#x03BC;m filter membrane, then filled into sterile glass bottles, and stored at 4&#x00B0;C in the dark for no more than 7&#x202F;days to prevent the introduction and proliferation of environmental microorganisms. We retained 30 sprouted plants in an illuminated incubator and kept them in light for 14&#x202F;h a day. On the 30th day after seeding emerged, the sweet clover plants were put into a light-proof beaker with 500&#x202F;mL ddH<sub>2</sub>O to clean them of nutrients. Then, incubation continued for 24&#x202F;h by using a 0.45&#x202F;&#x03BC;m water filter membrane to collect the root exudate solution, and through freeze drying, we obtained solid root exudates. Before use, the 0.45&#x202F;&#x03BC;m filter membrane was subjected to ultraviolet irradiation for 30&#x202F;min on both sides to ensure thorough sterilization. Subsequently, the tweezers were immersed in 75% ethanol for disinfection and briefly exposed to a flame for sterilization, followed by cooling to room temperature. These steps were taken to prevent the penetration or adhesion of fungal spores that could result from unsterilized materials or contamination during handling. This process was repeated until enough root exudates (the weight of the collected root system was greater than 300&#x202F;mg) were collected.</p>
<table-wrap position="float" id="tab1">
<label>Table 1</label>
<caption>
<p>Alpha of diversity indexes.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="top">No.</th>
<th align="center" valign="top">Ace</th>
<th align="center" valign="top">Chao</th>
<th align="center" valign="top">Shannon</th>
<th align="center" valign="top">Simpson (10<sup>&#x2212;2</sup>)</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="middle">C</td>
<td align="center" valign="top">1,569&#x202F;&#x00B1;&#x202F;43 d</td>
<td align="center" valign="top">1,576&#x202F;&#x00B1;&#x202F;61 d</td>
<td align="center" valign="top">5.08&#x202F;&#x00B1;&#x202F;0.05 c</td>
<td align="center" valign="top">1.31&#x202F;&#x00B1;&#x202F;0.02 d</td>
</tr>
<tr>
<td align="left" valign="middle">R1</td>
<td align="center" valign="top">1816&#x202F;&#x00B1;&#x202F;32 a</td>
<td align="center" valign="top">1810&#x202F;&#x00B1;&#x202F;55 a</td>
<td align="center" valign="top">5.51&#x202F;&#x00B1;&#x202F;0.11 b</td>
<td align="center" valign="top">1.63&#x202F;&#x00B1;&#x202F;0.01 a</td>
</tr>
<tr>
<td align="left" valign="middle">R2</td>
<td align="center" valign="top">1763&#x202F;&#x00B1;&#x202F;72 b</td>
<td align="center" valign="top">1757&#x202F;&#x00B1;&#x202F;47 b</td>
<td align="center" valign="top">5.57&#x202F;&#x00B1;&#x202F;0.05 a</td>
<td align="center" valign="top">1.42&#x202F;&#x00B1;&#x202F;0.01 c</td>
</tr>
<tr>
<td align="left" valign="middle">R3</td>
<td align="center" valign="top">1732&#x202F;&#x00B1;&#x202F;36 b</td>
<td align="center" valign="top">1732&#x202F;&#x00B1;&#x202F;44 b</td>
<td align="center" valign="top">5.58&#x202F;&#x00B1;&#x202F;0.07 a</td>
<td align="center" valign="top">1.52&#x202F;&#x00B1;&#x202F;0.01 b</td>
</tr>
<tr>
<td align="left" valign="middle">R4</td>
<td align="center" valign="top">1,668&#x202F;&#x00B1;&#x202F;19 c</td>
<td align="center" valign="top">1,685&#x202F;&#x00B1;&#x202F;13 c</td>
<td align="center" valign="top">5.55&#x202F;&#x00B1;&#x202F;0.07 b</td>
<td align="center" valign="top">1.41&#x202F;&#x00B1;&#x202F;0.03 c</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<p>&#x201C;Ace&#x201D; and &#x201C;Chao&#x201D; are indicators of species richness, &#x201C;Shannon&#x201D; and &#x201C;simpon&#x201D; are indicators of species evenness. The lowercase letters &#x201C;a, b, c, d&#x201D; in the table denote markers of statistical significance. Differences between data groups labeled with distinct letters are considered statistically significant (typically <italic>p</italic> &#x003C; 0.05).</p>
</table-wrap-foot>
</table-wrap>
<p>The main constituent substances of root exudates were determined by liquid chromatograph-mass spectrometer (LC&#x2013;MS), including 12 categories and 31 metabolites. The total carbon and nitrogen were determined by a TOC/TN analyzer. The TC was 64.2%, and the TN was 10.2% (<xref ref-type="bibr" rid="ref66">Xie et al., 2018</xref>; <xref ref-type="bibr" rid="ref34">Linlin et al., 2019</xref>).</p>
</sec>
<sec id="sec4">
<label>2.2</label>
<title>Field experiments</title>
<p>The field experiments were carried out in the waste dump of the Zhahanao&#x2019;er Open-Pit Coal Mine in Zhalute County, Inner Mongolia, China. The annual average temperature is 6.0&#x00B0;C, and the mean annual rainfall is approximately 379&#x202F;mm. Every test field block was covered with 20&#x202F;cm of the reconstructing soil, sized in 1&#x202F;m&#x202F;&#x00D7;&#x202F;1&#x202F;m. On 15 May 2023, <italic>Medicago sativa</italic> (<italic>Medicago Sativa Linn</italic>) was planted in blocks according to 150&#x202F;kg&#x202F;ha<sup>&#x2212;1</sup> seed. Five treatments and three replicates were designed: control (C, water) and four different root exudates&#x2019; application concentrations, as 40% mg/ml (R1), 55% mg/ml (R2), 70% mg/ml (R3), 85% mg/ml (R4), respectively, resulting in 15 field blocks in total.</p>
<p>In total, 100&#x202F;mL of root exudate solution was applied uniformly onto the soil when <italic>Medicago sativa</italic> was at seeding, branching, and flowering periods, respectively. During the entire plant life period, no fertilizer was added, and the plant was watered properly to keep the soil moist at all times. After 5&#x202F;months, we collected the 0&#x2013;20&#x202F;cm layer of soil with plants for analysis. Soil N-NH<sub>4</sub><sup>+</sup> and N-NO- 3 were determined by an AA3 analyzer (Technicon, Germany). Soil organic matter (OM) and TN were determined by an element analyzer (Elementar, Germany).</p>
</sec>
<sec id="sec5">
<label>2.3</label>
<title>DNA extraction and processing of pyrosequencing data</title>
<p>DNA was extracted from 0.5&#x202F;g of each individual replicate soil using a <italic>PowerSoil R DNA isolation kit</italic> (Mo Bio Laboratories, Inc., Carlsbad, CA, United States) (<xref ref-type="bibr" rid="ref69">Zhang et al., 2022</xref>). The hypervariable region V3&#x2013;V4 of the bacterial 16S rRNA gene was amplified with the primer pair 338F/806R (<xref ref-type="bibr" rid="ref67">Xu et al., 2016</xref>). The polymerase chain reaction (PCR) amplification method used <italic>TransGen AP221-02: TransStart FastPfu DNA Polymerase</italic> in a 20&#x202F;&#x03BC;L system.</p>
<p>DNA extraction, PCR amplification, fluorescence quantification, MiSeq library construction, and MiSeq sequencing were used to determine the community composition of soil bacteria. The production of PCR was detected by 2% of lipid gel in lipid sugar gel electrophoresis, and the library was constructed with the <italic>TruSeqTM DNA Sample Prep Kit</italic>. The qualified library was sequenced by the Silva Database (Release138<xref ref-type="fn" rid="fn0001"><sup>1</sup></xref>) on an <italic>Illumina MiSeq PE300</italic> platform (Illumina, San Diego, CA, United States) at <italic>Majorbio Bio-Pharm Technology Co., Ltd</italic>. (Shanghai, China).</p>
<p>The Raw Tags were PE Reads joined after splitting through <italic>FLASH</italic> and filtered by <italic>Qiime</italic> (Version 1.7.0) according to quality (<xref ref-type="bibr" rid="ref35">Lozupone et al., 2011</xref>). Finally, we found the effective tags through length filtering and chimera deletion (<xref ref-type="bibr" rid="ref4">Bokulich et al., 2013</xref>). Using a 97% identity threshold, the most abundant sequence from each operational taxonomic unit was selected as a representative sequence for that OTU (<xref ref-type="bibr" rid="ref33">Li et al., 2021</xref>). OTU clustering analysis and species taxonomy analysis were carried out based on the USEARCH7-uparse method.</p>
</sec>
<sec id="sec6">
<label>2.4</label>
<title>Statistical analyses</title>
<p>The alpha diversity indices were calculated using <italic>R</italic> (4.2.1; Simpson, Shannon, ACE, and Chao1, <xref ref-type="table" rid="tab1">Table 1</xref>). The beta diversity was performed by principal coordinates analysis (PCoA) and canonical correspondence analysis (CCA) based on <italic>UniFrac</italic> distance to compare the differences between the groups. <italic>R statistics analysis software</italic> and <italic>Origin statistics analysis software</italic> were carried out to calculate standard deviation, LSD, and Tukey&#x2019;s multiple range test through univariate statistical analysis and multivariate statistical analysis methods and to plot figures. The difference in means at a <italic>p-value of</italic> &#x003C; 0.05 was considered statistically significant.</p>
</sec>
</sec>
<sec sec-type="results" id="sec7">
<label>3</label>
<title>Results</title>
<sec id="sec8">
<label>3.1</label>
<title>Reconstituted soil fertility and plant growing</title>
<p>The concentration of soil TN and OM significantly increased with the increase in the root exudates&#x2019; amount (<xref ref-type="fig" rid="fig1">Figure 1</xref>). The N-NH<sub>4</sub><sup>+</sup> concentrations showed a trend of R2&#x202F;&#x003E;&#x202F;R3&#x202F;&#x003E;&#x202F;R1&#x202F;&#x003E;&#x202F;R4&#x202F;&#x003E;&#x202F;C, and the average growth rate of N-NH<sub>4</sub><sup>+</sup> was 11.3% (<xref ref-type="fig" rid="fig1">Figure 1a</xref>). The N-NO- 3 of R4 was the highest, corresponding to 20.8% higher than the control (<xref ref-type="fig" rid="fig1">Figure 1b</xref>). Compared with control, the root exudates&#x2019; application increased soil TN from 15.7 to 39.2% (<xref ref-type="fig" rid="fig1">Figure 1c</xref>). After using root exudates, reconstruction soil N-NH<sub>4</sub><sup>+</sup> and N-NO&#x2013; 3 increased significantly. There was no significant difference between the N-NO- 3 of R1, R2, and R3. The OM of R4 and R3 was higher than R1 (<xref ref-type="fig" rid="fig1">Figure 1d</xref>).</p>
<fig position="float" id="fig1">
<label>Figure 1</label>
<caption>
<p>Fertility of soil amended with root exudates. The letters &#x201C;a, b, c, d&#x201D; markers of statistical significance. Differences between data groups labeled with distinct letters are considered statistically significant (typically <italic>p</italic> &#x003C; 0.05).</p>
</caption>
<graphic xlink:href="fmicb-16-1466452-g001.tif">
<alt-text content-type="machine-generated">Four bar charts labeled a to d compare different soil parameters across treatments C, R1, R2, R3, and R4. Chart a shows N-NH4 levels, chart b shows N-NO3 levels, chart c shows TN levels, and chart d shows OM levels. Error bars indicate variability, and columns are marked with letters indicating statistically significant differences. C consistently has the lowest values, except in chart c.</alt-text>
</graphic>
</fig>
<p>The nourishing of root exudates made plant and root growth better than the control (<xref ref-type="fig" rid="fig2">Figure 2</xref>). The plant height and root length of R2 were the highest. The plant height of R1, R3, and R4 was similar, while the root length showed a trend of R3&#x202F;&#x003E;&#x202F;R4&#x202F;&#x003E;&#x202F;R1. The root-shoot ratio was significantly larger for root exudate treatments than for the control. Thus, the application of root exudates helped the plant root to grow a stem. Furthermore, treated plants&#x2019; root surface (<xref ref-type="fig" rid="fig2">Figure 2d</xref>), volume (<xref ref-type="fig" rid="fig2">Figure 2g</xref>), diameter (<xref ref-type="fig" rid="fig2">Figure 2f</xref>), number of root tips (<xref ref-type="fig" rid="fig2">Figure 2h</xref>), and branches (<xref ref-type="fig" rid="fig2">Figure 2e</xref>) were all higher than those of the control (R2 and R3 were always higher than the other). The correlation between the amount of root exudates applied and plant root growth: the root surface area, number of root tips, and average root diameter increased with the increase in the root exudates&#x2019; amount (<xref ref-type="fig" rid="fig3">Figure 3</xref>). Thus, the root growth resulted in increasing plant biomass (<xref ref-type="fig" rid="fig2">Figure 2i</xref>). The biomass and the root length followed the trend R2&#x202F;&#x003E;&#x202F;R3&#x202F;&#x003E;&#x202F;R4&#x202F;&#x003E;&#x202F;R1&#x202F;&#x003E;&#x202F;C.</p>
<fig position="float" id="fig2">
<label>Figure 2</label>
<caption>
<p>Plant root and biomass growth. The letters &#x201C;a, b, c, d&#x201D; markers of statistical significance. Differences between data groups labeled with distinct letters are considered statistically significant (typically <italic>p</italic> &#x003C; 0.05).</p>
</caption>
<graphic xlink:href="fmicb-16-1466452-g002.tif">
<alt-text content-type="machine-generated">Nine bar graphs labeled a to i depict various plant growth metrics across treatments C, R1, R2, R3, and R4. Graph (a) shows plant height, (b) root length, (c) root-to-shoot ratio, (d) root surface area, (e) number of root branches, (f) average root diameter, (g) root volume, (h) number of root tips, and (i) biomass. Each bar is marked with different letters indicating statistical significance, with measurements in centimeters, millimeters, or grams per day.</alt-text>
</graphic>
</fig>
<fig position="float" id="fig3">
<label>Figure 3</label>
<caption>
<p>Correlation between the amount of root exudates applied and plant root growth.</p>
</caption>
<graphic xlink:href="fmicb-16-1466452-g003.tif">
<alt-text content-type="machine-generated">Line graph showing the correlation between concentration and root characteristics: root surface area (red squares), number of root tips (yellow triangles), and average root diameter (blue circles). Equations and R-squared values indicate trends for each characteristic, with root surface area increasing, number of root tips peaking then declining, and average root diameter remaining stable.</alt-text>
</graphic>
</fig>
<p>The pH of the treated soil was 7.45, and there was no difference between the different amendments.</p>
</sec>
<sec id="sec9">
<label>3.2</label>
<title>Soil bacteria community composition</title>
<p><italic>Actinobacteriota</italic> and <italic>Proteobacteria</italic> were the dominant bacterial species in the amended soil. The sum of their relative abundance in the soil samples of each treatment was more than 50% (<xref ref-type="fig" rid="fig4">Figure 4</xref>). <italic>Chloroflexi</italic> and <italic>Firmicutes</italic> were the third and fourth largest relative type, respectively. The other bacteria types did not exceed 15%.</p>
<fig position="float" id="fig4">
<label>Figure 4</label>
<caption>
<p>Soil bacteria relative abundance of the phylum. &#x201C;p_&#x201D; is the abbreviation of &#x201C;Phylum&#x201D;.</p>
</caption>
<graphic xlink:href="fmicb-16-1466452-g004.tif">
<alt-text content-type="machine-generated">Stacked bar chart comparing bacterial composition across five samples labeled C, R1, R2, R3, and R4. Dominant phyla include Actinobacteriota and Proteobacteria. Other phyla such as Chloroflexi, Firmicutes, and Bacteroidota are present in smaller proportions. A color-coded legend identifies phyla.</alt-text>
</graphic>
</fig>
<p>After root exudates&#x2019; application, the <italic>Actinobacteriota</italic> increased significantly, especially in R4, which was 15% higher than in the control (<xref ref-type="fig" rid="fig4">Figure 4</xref>). It can be seen that the root exudates significantly promoted the relative abundance of the <italic>Actinobacteria</italic> class (<xref ref-type="fig" rid="fig5">Figure 5a</xref>) and the <italic>Euzebyales</italic> order (<xref ref-type="fig" rid="fig6">Figure 6a</xref>). The classes <italic>Acidimicrobiia</italic> and <italic>Thermoleophilia</italic> were the other main bacterial classes (<xref ref-type="fig" rid="fig6">Figures 6b</xref>,<xref ref-type="fig" rid="fig6">c</xref>). The <italic>Actinomarinales</italic> order of <italic>Acidimicrobiia</italic> class in the R2 treatment was higher than others. The <italic>IMCC26256</italic> and <italic>Microtrichales</italic> orders of the <italic>Acidimicrobiia</italic> class, as well as the <italic>Solirubrobacterales</italic> and <italic>Gaiellales</italic> orders of the <italic>Thermoleophilia</italic> class, were not different among the different root exudates&#x2019; amount treatments.</p>
<fig position="float" id="fig5">
<label>Figure 5</label>
<caption>
<p>Relative abundance of soil Actinobacteriota <bold>(a)</bold>, Proteobacteria <bold>(b)</bold>, and Chloroflexi <bold>(c)</bold>. &#x201C;p_&#x201D; is the abbreviation of &#x201C;Phylum,&#x201D; &#x201C;c_&#x201D; is the abbreviation of &#x201C;Class&#x201D;.</p>
</caption>
<graphic xlink:href="fmicb-16-1466452-g005.tif">
<alt-text content-type="machine-generated">Three bar charts show the relative abundance of microbial classes within different phyla across samples C, R1, R2, R3, and R4. Chart a represents phylum Actinobacteriota with classes including Thermoleophilia and Acidimicrobiia. Chart b shows phylum Proteobacteria with classes Gammaproteobacteria and Alphaproteobacteria. Chart c depicts phylum Chloroflexi with multiple classes like Dehalococcoidia and Anaerolineae. Each chart includes a legend detailing the color coding for various microbial classes, along with a table displaying specific abundance values for each class in the samples.</alt-text>
</graphic>
</fig>
<fig position="float" id="fig6">
<label>Figure 6</label>
<caption>
<p>Relative abundance of soil Actinobacteria <bold>(a)</bold>, Acidimicrobiia <bold>(b)</bold>, Thermoleophilia <bold>(c)</bold>, Alphaproteobacteria <bold>(d)</bold>, Gammaproteobacteria <bold>(e)</bold>, Chloroflexia <bold>(f)</bold>, and Anaerolineae <bold>(g)</bold>. &#x201C;c_&#x201D; is the abbreviation of &#x201C;Class&#x201D; and &#x201C;o_&#x201D; is the abbreviation of &#x201C;Order&#x201D;.</p>
</caption>
<graphic xlink:href="fmicb-16-1466452-g006.tif">
<alt-text content-type="machine-generated">Seven bar charts display microbial community compositions across different classifications: Actinobacteria, Actinimicrobia, Thermoleophilia, Alphaproteobacteria, Gammaproteobacteria, Chloroflexia, and Anaerolineae. Each chart compares the relative abundance of various orders within these classes across five sample points labeled C, R1, R2, R3, and R4. Color-coded legends indicate specific orders, providing numerical values for relative abundance in corresponding tables.</alt-text>
</graphic>
</fig>
<p>In contrast, the relative abundance of <italic>Proteobacteria</italic> was the highest under the control treatment, and lowest under the R4 treatment (<xref ref-type="fig" rid="fig4">Figure 4</xref>). The main <italic>Proteobacteria</italic> were the <italic>Alphaproteobacteria</italic> and <italic>Gammaproteobacteria</italic> (<xref ref-type="fig" rid="fig5">Figures 5b</xref>, <xref ref-type="fig" rid="fig6">6e</xref>) (<xref ref-type="fig" rid="fig5">Figure 5b</xref>). There was no difference among treatments for the <italic>Alphaproteobacteria</italic> class (<xref ref-type="fig" rid="fig6">Figure 6d</xref>). The exudates amendment of R1 reduced the <italic>1,013-28-CG33</italic> order but increased the <italic>Alteromonadales</italic> order. After root exudates were added, the number of orders under <italic>Gammaproteobacteria</italic> increased, and the increased bacterial orders were <italic>Alteromonadales</italic>, <italic>Burkholderiales</italic>, <italic>CCD24,</italic> and <italic>Cellvibrionales</italic>. The relative abundance significantly decreased for <italic>Alteromonadales</italic> with the addition of root exudates.</p>
<p>The <italic>Chloroflexi</italic> relative abundance after root exudates&#x2019; amendment increased slightly compared with the control (<xref ref-type="fig" rid="fig4">Figure 4</xref>). <italic>Chloroflexia</italic> and <italic>Anaerolineae</italic> were the main bacterial classes of the <italic>Chloroflexi</italic> phylum (<xref ref-type="fig" rid="fig5">Figure 5c</xref>). Further analysis for the orders of <italic>Thermomicrobiales</italic>, <italic>SBR1031,</italic> and <italic>Caldilineales</italic> showed the different amendment did not alter the relative abundance and community (<xref ref-type="fig" rid="fig6">Figures 6f</xref>,<xref ref-type="fig" rid="fig6">g</xref>). This indicates that root exudates helped the bacterial species <italic>Chloroflexia</italic> growth.</p>
</sec>
<sec id="sec10">
<label>3.3</label>
<title>Bacterial diversity and abundance</title>
<p>The different root exudates&#x2019; amendments and control all had a coverage rate of over 99% of the bacterial community in the tests. According to the alpha diversity results, root exudates&#x2019; amendment increased the alpha diversity indices (<xref ref-type="fig" rid="fig7">Figure 7</xref>). The Ace and Chao index of R1 was the highest, and then R2, R3, and R4. R2 and R3 had the highest Shannon index, followed by R1 and R4. However, the Simpson index showed a trend of C&#x202F;&#x003C;&#x202F;R4, R2&#x202F;&#x003C;&#x202F;R3&#x202F;&#x003C;&#x202F;R1. According to the correlation analyses (<xref ref-type="fig" rid="fig7">Figure 7</xref>), the alpha diversity was linearly correlated with root exudate concentration. To summarize the alpha diversity results, root exudates&#x2019; amendment tends to increase the homogeneity of developed soil bacteria species. It promotes local existing soil bacterial growth but does not stimulate new bacterial species. Thus, the number of soil bacteria increases rather than enhancing the species number.</p>
<fig position="float" id="fig7">
<label>Figure 7</label>
<caption>
<p>Correlation between the amount of applied root exudates and alpha bacteria diversity. &#x201C;Ace&#x201D; and &#x201C;Chao&#x201D; are indicators of species richness, &#x201C;Shannon&#x201D; and &#x201C;simpon&#x201D; are indicators of species evenness.</p>
</caption>
<graphic xlink:href="fmicb-16-1466452-g007.tif">
<alt-text content-type="machine-generated">Four scatter plots depict relationships between root exudates percentages and microbial diversity indices. Plots are labeled a to d. Each plot shows a linear regression line with respective equations and R-squared values: a) Ace index, b) Chao index, c) Shannon index, d) Simpson index. Data points are marked C for control and R1-R4 for different treatment levels. Error bars indicate variability.</alt-text>
</graphic>
</fig>
<p>The PCoA analysis evaluated the between-habitat diversity of amended soil bacteria (<xref ref-type="fig" rid="fig8">Figure 8</xref>). The first two principal components, PC1 and PC2, together explain 95.9% of the total variation in the bacterial community. The treatments by root exudates were significantly divided. R1, R2, and R3 gathered more closely than R4. The reason for this phenomenon might be that the content of root exudates has differential effects on bacterial diversity. When the concentration of root exudates ranges from 40 to 70%, bacterial diversity is at its highest. However, when the concentration of root exudates is 85%, the addition of root exudates leads to an imbalance in the utilization of soil nutrients by bacteria, thereby reducing bacterial diversity and causing a decrease in bacterial abundance.</p>
<fig position="float" id="fig8">
<label>Figure 8</label>
<caption>
<p>PCoA analysis of amended soil bacterial communities.</p>
</caption>
<graphic xlink:href="fmicb-16-1466452-g008.tif">
<alt-text content-type="machine-generated">Scatter plot showing principal component analysis with PC1 accounting for 90.1% and PC2 for 5.8% of the variance. Different colored dots represent categories C, R1, R2, R3, and R4, forming clusters.</alt-text>
</graphic>
</fig>
</sec>
<sec id="sec11">
<label>3.4</label>
<title>Relationships between amended soil bacterial communities</title>
<p>Canonical correspondence analysis was used to reveal what environmental factors changed the bacterial communities in the amended soil (<xref ref-type="fig" rid="fig9">Figure 9</xref>). The five treatments resulted in different bacterial distributions. The control was far away from amendments. The root length and plant height were positively associated with amended soil TN and OM. The shifts in bacteria OTUs were correlated with environmental variables. The root exudates changed the amended soil bacteria community by increasing plant root growth and soil fertility from increased TN and OM. R2 and R3 were significantly different from R1 and R4 in terms of biomass and fertility, in which R1 has the strongest promoting effect and R4 the weakest. The root exudates, thus, accelerate the transfer of soil nutrition to soil microorganisms.</p>
<fig position="float" id="fig9">
<label>Figure 9</label>
<caption>
<p>CCA analysis of amended soil bacterial communities.</p>
</caption>
<graphic xlink:href="fmicb-16-1466452-g009.tif">
<alt-text content-type="machine-generated">Scatter plot with CCA1 and CCA2 axes, accounting for 60.02% and 13.55% variance, respectively. Different colored circles represent groups C, R1, R2, R3, and R4, with a p-value of 0.01. Blue arrows indicate variables: root length, plant height, biomass, TN, and OM.</alt-text>
</graphic>
</fig>
</sec>
</sec>
<sec sec-type="discussion" id="sec12">
<label>4</label>
<title>Discussion</title>
<p>A main ecological problem from open coal mine use is restoring the soil surface environment. This is due to the shortage of topsoil and low soil fertility in many arid and semi-arid regions. Soil improvement is a critical component of ecological restoration in open-pit mining areas (<xref ref-type="bibr" rid="ref6">Cant&#x00F3; et al., 2020</xref>; <xref ref-type="bibr" rid="ref38">Malik and Jawad, 2022</xref>). Traditional fertilizers and root exudates, as two primary soil improvement methods, exhibit significant differences in their mechanisms and effects. Traditional fertilizers include chemical and organic fertilizers. The mechanism of chemical fertilizer is to improve soil fertility by directly supplementing nitrogen, phosphorus, potassium, and other nutrients in the soil (<xref ref-type="bibr" rid="ref54">Tong et al., 2024</xref>). Studies have shown that chemical fertilizers can significantly increase the content of available nutrients in soil; for example, nitrogen fertilizers can raise soil alkali-hydrolyzable nitrogen content by 30&#x2013;50% (<xref ref-type="bibr" rid="ref10">Dakora and Phillips, 2002</xref>; <xref ref-type="bibr" rid="ref2">Badri and Vivanco, 2009</xref>). However, the long-term application of chemical fertilizers may lead to adverse effects, such as soil acidification, compaction, and reduced microbial diversity (<xref ref-type="bibr" rid="ref9005">Wan et al., 2018</xref>).</p>
<p>The mechanism of action of organic fertilizer is to convert organic matter into humus through microbial decomposition, thereby improving soil structure and enhancing its water-holding and nutrient retention capacity (<xref ref-type="bibr" rid="ref62">Wen et al., 2020</xref>). Research shows that the application of organic fertilizer can significantly increase soil organic matter content. For instance, applying cow manure or biogas slurry can increase the soil organic matter content by more than 30% (<xref ref-type="bibr" rid="ref18">Hammad et al., 2020</xref>). Moreover, organic fertilizer can promote the formation of soil aggregates, improve soil aeration, and enhance the water&#x2013;air ratio (<xref ref-type="bibr" rid="ref9007">Wang et al., 2024</xref>). However, the improvement effect of organic fertilizer is relatively slow, and its contribution to enhancing soil microbial diversity is limited (<xref ref-type="bibr" rid="ref53">Tittonell et al., 2008</xref>). Root exudates indirectly improve soil structure and quality by activating insoluble nutrients and promoting microbial activity. Organic acids (e.g., oxalic acid and citric acid) in root exudates facilitate the dissolution of insoluble minerals through acidolysis and chelation (<xref ref-type="bibr" rid="ref3">Bais et al., 2006</xref>). For instance, root exudates from cypress can increase soil available phosphorus content by 20&#x2013;30%. Additionally, root exudates provide abundant microbial nutrients and carbon sources, enhancing rhizosphere microbial metabolic activity and accelerating nutrient mineralization and transformation (<xref ref-type="bibr" rid="ref31">Li et al., 2012</xref>; <xref ref-type="bibr" rid="ref14">Deru et al., 2023</xref>). Polysaccharides in root exudates also promote the formation of soil aggregates, thereby improving soil structure (<xref ref-type="bibr" rid="ref68">Zeng et al., 2008</xref>; <xref ref-type="bibr" rid="ref9008">Yang et al., 2024</xref>). Through literature review and data comparison (<xref ref-type="bibr" rid="ref20">Hao et al., 2022</xref>; <xref ref-type="bibr" rid="ref42">Nannipieri et al., 2008</xref>; <xref ref-type="bibr" rid="ref9006">Shen et al., 2019</xref>), the soil improvement effect indicators of traditional fertilization and applied root secretion were quantitatively analyzed and compared, as shown in <xref ref-type="table" rid="tab2">Table 2</xref>.</p>
<table-wrap position="float" id="tab2">
<label>Table 2</label>
<caption>
<p>Comparison of the effect data of different methods.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="top">Index</th>
<th align="left" valign="top">Chemical fertilizer</th>
<th align="left" valign="top">Organic fertilizer</th>
<th align="left" valign="top">Root exudates</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="top">Alkali-hydrolyzed nitrogen</td>
<td align="left" valign="top">Increased by 30&#x2013;50%</td>
<td align="left" valign="top">Increased by more than 30%</td>
<td align="left" valign="top">Increased by 10&#x2013;20%</td>
</tr>
<tr>
<td align="left" valign="top">Available phosphorus</td>
<td align="left" valign="top">Increased by 20&#x2013;40%</td>
<td align="left" valign="top">Increased by 20&#x2013;40%</td>
<td align="left" valign="top">Increased by 20&#x2013;30%</td>
</tr>
<tr>
<td align="left" valign="top">Available potassium</td>
<td align="left" valign="top">Increased by 25&#x2013;35%</td>
<td align="left" valign="top">Increased by 25&#x2013;35%</td>
<td align="left" valign="top">Increased by 15&#x2013;25%</td>
</tr>
<tr>
<td align="left" valign="top">Microbial diversity</td>
<td align="left" valign="top">May reduce</td>
<td align="left" valign="top">Shannon index:<break/>Increased by 5&#x2013;10%;<break/>Number of OTUs:<break/>Increased by 10&#x2013;15%</td>
<td align="left" valign="top">Shannon index: Increased by 15&#x2013;25%;<break/>Number of OTUs: Adding 20&#x2013;30%</td>
</tr>
<tr>
<td align="left" valign="top">Structure improvement</td>
<td align="left" valign="top">No significant improvement</td>
<td align="left" valign="top">Aggregate stability:<break/>Increased by 10&#x2013;20%;<break/>Porosity:<break/>No significant change;<break/>Proportion of water-stable aggregates:<break/>Increased by 5&#x2013;10%</td>
<td align="left" valign="top">Aggregate stability:<break/>Increased by 30&#x2013;40%;<break/>Porosity:<break/>Increased by 15&#x2013;25%;<break/>Proportion of water-stable aggregates:<break/>Increased by 20&#x2013;30%</td>
</tr>
</tbody>
</table>
</table-wrap>
<p>In summary, although traditional fertilization has relatively better immediate effects on improving soil nutrients, considering the flexibility and sustainability of the improvement effect, the improved root secretion method is more suitable for open-pit soil with poor nutrients, high soil strength, and low microbial activity (<xref ref-type="table" rid="tab2">Table 2</xref>) (<xref ref-type="bibr" rid="ref13">Deonalli et al., 2017</xref>; <xref ref-type="bibr" rid="ref71">Zhao et al., 2025</xref>; <xref ref-type="bibr" rid="ref28">Keiluweit et al., 2015</xref>). In this study, root exudates were used as soil amendments to improve the soil content of C and N in an abandoned open-pit coal mine area. Soil OM is the limiting factor to the growth of both plants and microbes (<xref ref-type="bibr" rid="ref29">Lal, 2020</xref>). The root exudates contain vitamins, peptides, organic acids, lipids, carbohydrates, and other important plant organic nutrient substances. These substances promote root growth, absorb more water from the soil, and support the whole plant growing stage (<xref ref-type="bibr" rid="ref19">Han et al., 2018</xref>). Furthermore, secondary metabolites in root exudates can help plants adapt to poor fertility and the biotic stress of impoverished soils (<xref ref-type="bibr" rid="ref56">Vives-Peris et al., 2020</xref>). The results of this study also showed that the higher the amount of root exudates, the higher the soil fertility and root-shoot ratio.</p>
<p>Microorganisms play an important role in the nutrient transformation between amended soil and plants. However, the microorganism density is low with small diversity, especially in the mine ecosystem (<xref ref-type="bibr" rid="ref73">Zhou et al., 2018</xref>). Previous studies have shown that root exudates can stimulate the growth of microorganisms and improve soil bacterial abundance (<xref ref-type="bibr" rid="ref72">Zhou et al., 2012</xref>). When root exudates are sufficient, they can provide more energy for microbial metabolism and improve the abundance of soil bacteria, which is consistent with the conclusion of this study. On the one hand, the improved fertility of amended soil provides a substrate that is easily decomposed by microorganisms (<xref ref-type="bibr" rid="ref8">Clocchiatti et al., 2021</xref>). The soil bacteria density increased with the continuous supply of nutrients (<xref ref-type="fig" rid="fig7">Figure 7</xref>). This was shown by the positive correlation between soil bacterial community distribution and soil fertility from OM and TN (<xref ref-type="fig" rid="fig9">Figure 9</xref>). Furthermore, increased plant growth by adding root exudates improved the ecological function of amended soil (<xref ref-type="bibr" rid="ref44">Parray et al., 2016</xref>). Plant growth promotes rhizobacteria interactions with plants, such as element fixation and solubilization of N and P (<xref ref-type="bibr" rid="ref49">Rosier et al., 2018</xref>), as well as improving plant drought resistance and the antiviral ability in the rhizosphere (<xref ref-type="bibr" rid="ref41">Morales et al., 2022</xref>). Thus, root exudates improve soil bacteria by promoting plant growth, especially root growth (<xref ref-type="bibr" rid="ref9002">Ge et al., 2017</xref>).</p>
<p>In our experiments, the local dominating bacteria community was driven by root exudates and plant growth, as also shown in previous studies (<xref ref-type="bibr" rid="ref16">Freedman et al., 2015</xref>; <xref ref-type="bibr" rid="ref30">Lanoue et al., 2010</xref>; <xref ref-type="bibr" rid="ref39">Maurer et al., 2021</xref>). The bacteria were sensitive to nutrients and organic matter in poor soils (<xref ref-type="bibr" rid="ref2">Badri and Vivanco, 2009</xref>). The results of this study indicate that the bacterial diversity was highest for a root exudate concentration of 40&#x2013;70%. If the added root exudates cause an imbalance between C and N in terms of bacterial use, the diversity of bacteria could be reduced (<xref ref-type="bibr" rid="ref25">Jiang et al., 2021</xref>). Consequently, bacterial abundance decreased for a root exudate concentration of 85%.</p>
<p>A high concentration of root exudates was most obvious in promoting <italic>Actinobacteriota</italic>, <italic>Proteobacteria,</italic> and <italic>Chloroflexi</italic>. <italic>Actinobacteriota</italic> are an important bacterium in soil. Its slender and complex mycelia contribute to the formation of soil aggregates and the accumulation of organic carbon (<xref ref-type="bibr" rid="ref65">Xie et al., 2022</xref>), which is of great significance to the improvement of basic soil fertility (<xref ref-type="bibr" rid="ref21">Hern&#x00E1;ndez-&#x00C1;lvarez et al., 2022</xref>). <italic>Proteobacteria</italic> are the most widely distributed and complex functional flora in farmland and natural soil. <italic>Proteobacteria</italic> help the fixation and transformation, especially mineralization, in the soil N cycle (<xref ref-type="bibr" rid="ref32">Li et al., 2022</xref>). <italic>Alphaproteobacteria</italic> and <italic>Gammaproteobacteria</italic> are closely related to plant N-uptake and utilization from soil (<xref ref-type="bibr" rid="ref5">Bugg et al., 2011</xref>). <italic>Chloroflexi</italic> is a facultative anaerobic bacteria that can generate energy through photosynthesis (<xref ref-type="bibr" rid="ref59">Wang et al., 2019</xref>). It does not produce oxygen or fix nitrogen in photosynthesis, but it is helpful for the circulation of soil nutrients (<xref ref-type="bibr" rid="ref50">Speirs et al., 2019</xref>). <italic>Firmicutes</italic> can produce spores with strong resistance to drought and extreme environments. All abundant bacteria in our experiments can adapt well to harsh soil conditions (<xref ref-type="bibr" rid="ref21">Hern&#x00E1;ndez-&#x00C1;lvarez et al., 2022</xref>). They, as well, can improve the efficiency of amended soil to reconstruct and enhance plant growth.</p>
<p>Activating soil nutrients through root exudates and promoting the absorption of soil nutrients by crops is of great significance for improving the productivity of poor soil. The soil in open-pit mining areas is nutrient-poor, making it essential to select locally adapted reclamation plants that are most suitable for growth in the planting system. Additionally, the impact of root exudates from these reclamation plants on soil properties and plant growth should be considered. Direct or indirect effects of root secretion from reclaimed plant species can affect the growth and development of their own or neighboring crops, improving the properties of soil under nutrient stress, and enhancing the effect of plant reclamation. The findings regarding the interaction between plant root exudates and rhizosphere microorganisms in this study hold significant theoretical and practical value for elucidating soil ecosystem functions and guiding vegetation restoration and ecological management in mined areas. In the future, the appropriate ecological planting model and adaptation mechanism of plants mediated by plant root exudates can be systematically studied to guide the ecological planting work in mines.</p>
</sec>
<sec sec-type="conclusions" id="sec13">
<label>5</label>
<title>Conclusion</title>
<p>Applying root exudates in impoverished soil ecosystems of open-pit coal mines could increase soil fertility and improve the plant nutrient status. In the amended soil, the root exudates can simultaneously promote plant growth and bacterial proliferation. The growing plant roots will gradually increase bacterial abundance and diversity. The local dominant bacterial species of soil nutrient cycling increased and became more abundant. In a certain range, increasing the amount of root exudates can significantly increase the bacterial abundance.</p>
<p>This study demonstrates the practical application feasibility of root exudate-mediated soil amendment technology in the ecological restoration of open-pit coal mines. However, prior to large-scale implementation, region-specific analyses integrating pedological characteristics, reclamation objectives, and monitoring technologies must be conducted. A three-phase implementation strategy should be rigorously followed. In the initial restoration stage, concentrated root exudate solutions should be sprayed preferentially to activate the local microbial community and create a micro-environment for subsequent plant colonization. In the mid-term collaborative stage, synchronized with the sowing of stress-tolerant plants, continuous input of low-concentration exudates from the drip irrigation system should be adopted to promote root development and colonization of microorganisms. In the later maintenance stage, soil microbial diversity should be regularly monitored, and the amount of exudate application should be adjusted according to the abundance of dominant bacteria to avoid excessive stimulation that leads to community imbalance.</p>
</sec>
</body>
<back>
<sec sec-type="data-availability" id="sec14">
<title>Data availability statement</title>
<p>The original contributions presented in the study are publicly available. This data can be found here: <ext-link xlink:href="https://data.mendeley.com/datasets/jm2gcrf66p/1" ext-link-type="uri">https://data.mendeley.com/datasets/jm2gcrf66p/1</ext-link>.</p>
</sec>
<sec sec-type="author-contributions" id="sec15">
<title>Author contributions</title>
<p>ZY: Funding acquisition, Resources, Writing &#x2013; review &#x0026; editing. JN: Supervision, Writing &#x2013; review &#x0026; editing. TW: Data curation, Visualization, Writing &#x2013; original draft. JL: Formal analysis, Writing &#x2013; original draft. LZ: Conceptualization, Writing &#x2013; review &#x0026; editing. XC: Investigation, Writing &#x2013; review &#x0026; editing. RB: Validation, Writing &#x2013; review &#x0026; editing.</p>
</sec>
<sec sec-type="funding-information" id="sec16">
<title>Funding</title>
<p>The author(s) declare that financial support was received for the research and/or publication of this article. This research was funded by Natural Science Foundation of Liaoning Province Grant No. 2023-MS-342.</p>
</sec>
<sec sec-type="COI-statement" id="sec17">
<title>Conflict of interest</title>
<p>ZY was employed by China Coal Technology &#x0026; Engineering Group Shenyang Engineering Company. The remaining 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="disclaimer" id="sec18">
<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="sec23" sec-type="supplementary-material">
<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.2025.1466452/full#supplementary-material" ext-link-type="uri">https://www.frontiersin.org/articles/10.3389/fmicb.2025.1466452/full#supplementary-material</ext-link></p>
<supplementary-material xlink:href="Data_Sheet_1.xlsx" id="SM1" mimetype="application/vnd.openxmlformats-officedocument.spreadsheetml.sheet" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</sec>
<fn-group>
<fn id="fn0001"><p><sup>1</sup><ext-link xlink:href="http://www.arb-silva.de" ext-link-type="uri">http://www.arb-silva.de</ext-link></p></fn>
</fn-group>
<ref-list>
<title>References</title>
<ref id="ref1"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Alberto</surname> <given-names>C.</given-names></name> <name><surname>Kaiser</surname> <given-names>C.</given-names></name> <name><surname>Merchant</surname> <given-names>A.</given-names></name> <name><surname>Richter</surname> <given-names>A.</given-names></name> <name><surname>Wanek</surname> <given-names>W</given-names></name></person-group>. (<year>2019</year>). <article-title>Root exudation of primary metabolites: mechanisms and their roles in plant responses to environmental stimuli</article-title>. <source>Front. Plant Sci.</source> <volume>10</volume>:<fpage>157</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fpls.2019.00157</pub-id></citation></ref>
<ref id="ref2"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Badri</surname> <given-names>D. V.</given-names></name> <name><surname>Vivanco</surname> <given-names>J. M.</given-names></name></person-group> (<year>2009</year>). <article-title>Regulation and function of root exudates</article-title>. <source>Plant Cell Environ.</source> <volume>32</volume>, <fpage>666</fpage>&#x2013;<lpage>681</lpage>. doi: <pub-id pub-id-type="doi">10.1111/j.1365-3040.2009.01926.x</pub-id>, PMID: <pub-id pub-id-type="pmid">19143988</pub-id></citation></ref>
<ref id="ref3"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Bais</surname> <given-names>H. P.</given-names></name> <name><surname>Weir</surname> <given-names>T. L.</given-names></name> <name><surname>Perry</surname> <given-names>L. G.</given-names></name> <name><surname>Gilroy</surname> <given-names>S.</given-names></name> <name><surname>Vivanco</surname> <given-names>J. M.</given-names></name></person-group> (<year>2006</year>). <article-title>The role of root exudates in rhizosphere interactions with plants and other organisms</article-title>. <source>Annu. Rev. Plant Biol.</source> <volume>57</volume>, <fpage>233</fpage>&#x2013;<lpage>266</lpage>. doi: <pub-id pub-id-type="doi">10.1146/annurev.arplant.57.032905.105159</pub-id>, PMID: <pub-id pub-id-type="pmid">16669762</pub-id></citation></ref>
<ref id="ref4"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Bokulich</surname> <given-names>N. A.</given-names></name> <name><surname>Subramanian</surname> <given-names>S.</given-names></name> <name><surname>Faith</surname> <given-names>J. J.</given-names></name> <name><surname>Gevers</surname> <given-names>D.</given-names></name> <name><surname>Gordon</surname> <given-names>J. I.</given-names></name> <name><surname>Knight</surname> <given-names>R.</given-names></name> <etal/></person-group>. (<year>2013</year>). <article-title>Quality-filtering vastly improves diversity estimates from Illumina amplicon sequencing</article-title>. <source>Nat. Methods</source> <volume>10</volume>, <fpage>57</fpage>&#x2013;<lpage>59</lpage>. doi: <pub-id pub-id-type="doi">10.1038/nmeth.2276</pub-id>, PMID: <pub-id pub-id-type="pmid">23202435</pub-id></citation></ref>
<ref id="ref5"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Bugg</surname> <given-names>T. D. H.</given-names></name> <name><surname>Ahmad</surname> <given-names>M.</given-names></name> <name><surname>Hardiman</surname> <given-names>E. M.</given-names></name> <name><surname>Singh</surname> <given-names>R.</given-names></name></person-group> (<year>2011</year>). <article-title>The emerging role for bacteria in lignin degradation and bio-product formation</article-title>. <source>Curr. Opin. Biotechnol.</source> <volume>22</volume>, <fpage>394</fpage>&#x2013;<lpage>400</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.copbio.2010.10.009</pub-id>, PMID: <pub-id pub-id-type="pmid">21071202</pub-id></citation></ref>
<ref id="ref6"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Cant&#x00F3;</surname> <given-names>C. D. L. F.</given-names></name> <name><surname>Simonin</surname> <given-names>M.</given-names></name> <name><surname>King</surname> <given-names>E.</given-names></name> <name><surname>Moulin</surname> <given-names>L.</given-names></name> <name><surname>Bennett</surname> <given-names>M. J.</given-names></name> <name><surname>Castrillo</surname> <given-names>G.</given-names></name> <etal/></person-group>. (<year>2020</year>). <article-title>An extended root phenotype: the rhizosphere, its formation and impacts on plant fitness</article-title>. <source>Plant J.</source> <volume>103</volume>, <fpage>951</fpage>&#x2013;<lpage>964</lpage>. doi: <pub-id pub-id-type="doi">10.1111/tpj.14781</pub-id>, PMID: <pub-id pub-id-type="pmid">32324287</pub-id></citation></ref>
<ref id="ref9001"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Carla</surname> <given-names>A.</given-names></name> <name><surname>Catherine</surname> <given-names>C.</given-names></name> <name><surname>H&#x00E9;l&#x00E8;ne</surname> <given-names>G.</given-names></name></person-group> (<year>2020</year>). <article-title>Challenges for cysteamine stabilization, quantification, and biological effects improvement</article-title>. <source>J. Pharm. Anal.</source> <volume>10</volume>, <fpage>499</fpage>&#x2013;<lpage>516</lpage>.</citation></ref>
<ref id="ref7"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Chen</surname> <given-names>B.</given-names></name> <name><surname>Zhang</surname> <given-names>Y.</given-names></name> <name><surname>Rafiq</surname> <given-names>M. T.</given-names></name> <name><surname>Khan</surname> <given-names>K. Y.</given-names></name> <name><surname>Pan</surname> <given-names>F.</given-names></name> <name><surname>Yang</surname> <given-names>X.</given-names></name> <etal/></person-group>. (<year>2014</year>). <article-title>Improvement of cadmium uptake and accumulation in Sedum alfredii by endophytic bacteria Sphingomonas SaMR12: effects on plant growth and root exudates</article-title>. <source>Chemosphere</source> <volume>117</volume>, <fpage>367</fpage>&#x2013;<lpage>373</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.chemosphere.2014.07.078</pub-id>, PMID: <pub-id pub-id-type="pmid">25169647</pub-id></citation></ref>
<ref id="ref8"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Clocchiatti</surname> <given-names>A.</given-names></name> <name><surname>Hannula</surname> <given-names>S. E.</given-names></name> <name><surname>van den Berg</surname> <given-names>M.</given-names></name> <name><surname>Hundscheid</surname> <given-names>M. P. J.</given-names></name> <name><surname>de Boer</surname> <given-names>W.</given-names></name></person-group> (<year>2021</year>). <article-title>Evaluation of phenolic root exudates as stimulants of Saptrophic Fungi in the rhizosphere</article-title>. <source>Front. Microbiol.</source> <volume>12</volume>:<fpage>644046</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fmicb.2021.644046</pub-id>, PMID: <pub-id pub-id-type="pmid">33936001</pub-id></citation></ref>
<ref id="ref9"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Cordovez</surname> <given-names>V.</given-names></name> <name><surname>Rotoni</surname> <given-names>C.</given-names></name> <name><surname>Dini-Andreote</surname> <given-names>F.</given-names></name> <name><surname>Oyserman</surname> <given-names>B.</given-names></name> <name><surname>Raaijmakers</surname> <given-names>J. M.</given-names></name></person-group> (<year>2024</year>). <article-title>Synergistic effects of rhizosphere effect and combined organic and chemical fertilizers application on soil bacterial diversity and community structure in oilseed rape cultivation</article-title>. <source>Front. Microbiol.</source> <volume>15</volume>:<fpage>1374199</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fmicb.2024.1374199</pub-id></citation></ref>
<ref id="ref10"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Dakora</surname> <given-names>F. D.</given-names></name> <name><surname>Phillips</surname> <given-names>D. A.</given-names></name></person-group> (<year>2002</year>). <article-title>Root exudates as mediators of mineral acquisition in low-nutrient environments</article-title>. <source>Plant Soil</source> <volume>245</volume>, <fpage>35</fpage>&#x2013;<lpage>47</lpage>. doi: <pub-id pub-id-type="doi">10.1023/A:1020809400075</pub-id></citation></ref>
<ref id="ref11"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Damin</surname> <given-names>V.</given-names></name> <name><surname>Franco</surname> <given-names>H. C. J.</given-names></name> <name><surname>Moraes</surname> <given-names>M. F.</given-names></name> <name><surname>Franco</surname> <given-names>A.</given-names></name> <name><surname>Trivelin</surname> <given-names>P. C. O.</given-names></name></person-group> (<year>2008</year>). <article-title>Nitrogen loss in <italic>Brachiaria decumbens</italic> after application of glyphosate or glufosinate-ammonium</article-title>. <source>Sci. Agr.</source> <volume>64</volume>, <fpage>402</fpage>&#x2013;<lpage>407</lpage>. doi: <pub-id pub-id-type="doi">10.1590/S0103-90162008000400012</pub-id></citation></ref>
<ref id="ref12"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Demyan</surname> <given-names>M. S.</given-names></name> <name><surname>Smeck</surname> <given-names>N.</given-names></name></person-group> (<year>2022</year>). <article-title>Chemical and physical temporal and spatial changes in 25-year-old mine soils in Southeast Ohio</article-title>. <source>Land Degard Dev</source> <volume>33</volume>, <fpage>33</fpage>&#x2013;<lpage>35</lpage>. doi: <pub-id pub-id-type="doi">10.1002/ldr.4150</pub-id></citation></ref>
<ref id="ref13"><citation citation-type="other"><person-group person-group-type="author"><name><surname>Deonalli</surname> <given-names>D.</given-names></name> <name><surname>Sharma</surname> <given-names>R.</given-names></name> <name><surname>Jangid</surname> <given-names>K.</given-names></name></person-group> (<year>2017</year>). &#x201C;<article-title>Microbial community dynamics during soil ecosystem development</article-title>&#x201D; in <source>Mining of microbial wealth and meta genomics</source>. <publisher-name>Springer</publisher-name>, <fpage>57</fpage>&#x2013;<lpage>82</lpage>. doi: <pub-id pub-id-type="doi">10.1007/978-981-10-5708-3_4</pub-id></citation></ref>
<ref id="ref14"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Deru</surname> <given-names>J. G. C.</given-names></name> <name><surname>Bloem</surname> <given-names>J.</given-names></name> <name><surname>de Goede</surname> <given-names>R.</given-names></name> <name><surname>Brussaard</surname> <given-names>L.</given-names></name> <name><surname>van Eekeren</surname> <given-names>N.</given-names></name></person-group> (<year>2023</year>). <article-title>Effects of organic and inorganic fertilizers on soil properties related to the regeneration of ecosystem services in peat grasslands</article-title>. <source>Appl. Soil Ecol.</source> <volume>187</volume>:<fpage>104838</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.apsoil.2023.104838</pub-id></citation></ref>
<ref id="ref15"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Feng</surname> <given-names>Z. R. L.</given-names></name></person-group> (<year>2019</year>). <article-title>Effects of surface coal mining and land reclamation on soil properties: a review</article-title>. <source>Earth Sci. Rev.</source> <volume>191</volume>, <fpage>12</fpage>&#x2013;<lpage>25</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.earscirev.2019.02.015</pub-id></citation></ref>
<ref id="ref16"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Freedman</surname> <given-names>Z. B.</given-names></name> <name><surname>Romanowicz</surname> <given-names>K. J.</given-names></name> <name><surname>Upchurch</surname> <given-names>R. A.</given-names></name> <name><surname>Zak</surname> <given-names>D. R.</given-names></name></person-group> (<year>2015</year>). <article-title>Differential responses of total and active soil microbial communities to long-term experimental N deposition</article-title>. <source>Soil Biol. Biochem.</source> <volume>90</volume>, <fpage>275</fpage>&#x2013;<lpage>282</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.soilbio.2015.08.014</pub-id></citation></ref>
<ref id="ref9002"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Ge</surname> <given-names>S. Y.</given-names></name> <name><surname>Zhou</surname> <given-names>H.</given-names></name> <name><surname>Song</surname> <given-names>J.</given-names></name> <name><surname>Yu</surname> <given-names>J.</given-names></name> <name><surname>Wei</surname> <given-names>W.</given-names></name> <name><surname>Li</surname> <given-names>S. Y.</given-names></name> <etal/></person-group>. (<year>2017</year>). <article-title>Interaction between Humic Acid and p-Arsanilic Acid Using Fluorescence Quenching Approach</article-title>. <source>Asian. J. Ecotoxic.</source> <volume>12</volume>, <fpage>667</fpage>&#x2013;<lpage>674</lpage>. doi: <pub-id pub-id-type="doi">10.7524/AJE.1673-5897.20161227002</pub-id></citation></ref>
<ref id="ref17"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Haichar</surname> <given-names>F. E.</given-names></name> <name><surname>Heulin</surname> <given-names>T.</given-names></name> <name><surname>Guyonnet</surname> <given-names>J. P.</given-names></name> <name><surname>Achouak</surname> <given-names>W.</given-names></name></person-group> (<year>2016</year>). <article-title>Stable isotope probing of carbon flow in the plant holobiont</article-title>. <source>Curr. Opin. Biotechnol.</source> <volume>41</volume>, <fpage>9</fpage>&#x2013;<lpage>13</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.copbio.2016.02.023</pub-id></citation></ref>
<ref id="ref18"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Hammad</surname> <given-names>H. M.</given-names></name> <name><surname>Khaliq</surname> <given-names>A.</given-names></name> <name><surname>Abbas</surname> <given-names>F.</given-names></name> <name><surname>Farhad</surname> <given-names>W.</given-names></name> <name><surname>Bakhat</surname> <given-names>H. F.</given-names></name></person-group> (<year>2020</year>). <article-title>Comparative effects of organic and inorganic fertilizers on soil organic carbon and wheat productivity under arid region</article-title>. <source>Commun. Soil Sci. Plant Anal.</source> <volume>51</volume>, <fpage>1406</fpage>&#x2013;<lpage>1422</lpage>. doi: <pub-id pub-id-type="doi">10.1080/00103624.2020.1763385</pub-id></citation></ref>
<ref id="ref9003"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Hamid</surname> <given-names>G. S.</given-names></name> <name><surname>Mikhail</surname> <given-names>V.</given-names></name> <name><surname>Tero-Petri</surname> <given-names>R.</given-names></name> <name><surname>Viktor</surname> <given-names>G.</given-names></name> <name><surname>Krzysztof</surname> <given-names>K.</given-names></name> <name><surname>&#x0141;apkowski</surname> <given-names>M</given-names></name></person-group>. (<year>2022</year>). <article-title>Towards electrochemical hydrogen storage in liquid organic hydrogen carriers via proton-coupled electron transfers</article-title>. <source>J. Energy Chem.</source> <volume>73</volume>, <fpage>292</fpage>&#x2013;<lpage>300</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.jechem.2022.06.015</pub-id></citation></ref>
<ref id="ref19"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Han</surname> <given-names>X.</given-names></name> <name><surname>Xu</surname> <given-names>C.</given-names></name> <name><surname>Dungait</surname> <given-names>J. A. J.</given-names></name> <name><surname>Bol</surname> <given-names>R.</given-names></name> <name><surname>Wang</surname> <given-names>X.</given-names></name> <name><surname>Wu</surname> <given-names>W.</given-names></name> <etal/></person-group>. (<year>2018</year>). <article-title>Straw incorporation increases crop yield and soil organic carbon sequestration but varies under different natural conditions and farming practices in China: a system analysis</article-title>. <source>Biogeosciences</source> <volume>15</volume>, <fpage>1933</fpage>&#x2013;<lpage>1946</lpage>. doi: <pub-id pub-id-type="doi">10.5194/bg-15-1933-2018</pub-id></citation></ref>
<ref id="ref20"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Hao</surname> <given-names>C.</given-names></name> <name><surname>Dungait</surname> <given-names>J. A. J.</given-names></name> <name><surname>Wei</surname> <given-names>X.</given-names></name> <name><surname>Ge</surname> <given-names>T.</given-names></name> <name><surname>Kuzyakov</surname> <given-names>Y.</given-names></name> <name><surname>Cui</surname> <given-names>Z.</given-names></name> <etal/></person-group>. (<year>2022</year>). <article-title>Maize root exudate composition alters rhizosphere bacterial community to control hotspots of hydrolase activity in response to nitrogen supply</article-title>. <source>Soil Biol. Biochem.</source> <volume>170</volume>:<fpage>108717</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.soilbio.2022.108717</pub-id></citation></ref>
<ref id="ref21"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Hern&#x00E1;ndez-&#x00C1;lvarez</surname> <given-names>C.</given-names></name> <name><surname>Garc&#x00ED;a-Oliva</surname> <given-names>F.</given-names></name> <name><surname>Cruz-Ortega</surname> <given-names>R.</given-names></name></person-group> (<year>2022</year>). <article-title>Squash root microbiome transplants and metagenomic inspection for in situ arid adaptations</article-title>. <source>Sci. Total Environ.</source> <volume>805</volume>:<fpage>150136</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.scitotenv.2021.150136</pub-id>, PMID: <pub-id pub-id-type="pmid">34818799</pub-id></citation></ref>
<ref id="ref22"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Hu</surname> <given-names>A.</given-names></name> <name><surname>Xu</surname> <given-names>S.</given-names></name> <name><surname>Qin</surname> <given-names>D.</given-names></name> <name><surname>Li</surname> <given-names>W.</given-names></name> <name><surname>Zhao</surname> <given-names>X.</given-names></name></person-group> (<year>2021</year>). <article-title>Role of silicon in mediating phosphorus imbalance in plants</article-title>. <source>Plan. Theory</source> <volume>10</volume>:<fpage>51</fpage>. doi: <pub-id pub-id-type="doi">10.3390/plants10010051</pub-id></citation></ref>
<ref id="ref23"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Huang</surname> <given-names>A. C.</given-names></name> <name><surname>Jiang</surname> <given-names>T.</given-names></name> <name><surname>Liu</surname> <given-names>Y. X.</given-names></name> <name><surname>Bai</surname> <given-names>Y. C.</given-names></name> <name><surname>Reed</surname> <given-names>J.</given-names></name> <name><surname>Qu</surname> <given-names>B.</given-names></name> <etal/></person-group>. (<year>2019</year>). <article-title>A specialized metabolic network selectively modulates Arabidopsis root microbiota</article-title>. <source>Science</source> <volume>364</volume>:<fpage>546</fpage>. doi: <pub-id pub-id-type="doi">10.1126/science.aau6389</pub-id>, PMID: <pub-id pub-id-type="pmid">31073042</pub-id></citation></ref>
<ref id="ref24"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Jiang</surname> <given-names>H.</given-names></name> <name><surname>Halverson</surname> <given-names>L. J.</given-names></name> <name><surname>Dong</surname> <given-names>L.</given-names></name></person-group> (<year>2022</year>). <article-title>A miniaturized bioelectricity generation device using plant root exudates to feed electrogenic bacteria</article-title>. <source>Sensor. Actuat. A-Phys.</source> <volume>342</volume>:<fpage>113649</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.sna.2022.113649</pub-id></citation></ref>
<ref id="ref25"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Jiang</surname> <given-names>Z.</given-names></name> <name><surname>Liu</surname> <given-names>Y.</given-names></name> <name><surname>Yang</surname> <given-names>J.</given-names></name> <name><surname>Zhou</surname> <given-names>Z.</given-names></name> <name><surname>Gunina</surname> <given-names>A.</given-names></name></person-group> (<year>2021</year>). <article-title>Effects of nitrogen fertilization on the rhizosphere priming</article-title>. <source>Plant Soil</source> <volume>462</volume>, <fpage>489</fpage>&#x2013;<lpage>503</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s11104-021-04872-6</pub-id></citation></ref>
<ref id="ref26"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Jones</surname> <given-names>D. L.</given-names></name> <name><surname>Yajun</surname> <given-names>J.</given-names></name> <name><surname>Tida</surname> <given-names>W.</given-names></name> <etal/></person-group>. (<year>2017</year>). <article-title>Stability and dynamics of enzyme activity patterns in the rice rhizosphere: effects of plant growth and temperature</article-title>. <source>Soil Biol. Biochem.</source> <volume>113</volume>, <fpage>108</fpage>&#x2013;<lpage>115</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.soilbio.2017.06.005</pub-id></citation></ref>
<ref id="ref27"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Ju</surname> <given-names>T.</given-names></name> <name><surname>Han</surname> <given-names>S.</given-names></name> <name><surname>Meng</surname> <given-names>Y.</given-names></name> <name><surname>Jiang</surname> <given-names>J.</given-names></name></person-group> (<year>2021</year>). <article-title>High-end reclamation of coal fly ash focusing on elemental extraction and synthesis of porous materials</article-title>. <source>ACS Sustain. Chem. Eng.</source> <volume>9</volume>, <fpage>6894</fpage>&#x2013;<lpage>6911</lpage>. doi: <pub-id pub-id-type="doi">10.1021/acssuschemeng.1c00587</pub-id></citation></ref>
<ref id="ref28"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Keiluweit</surname> <given-names>M.</given-names></name> <name><surname>Bougoure</surname> <given-names>J. J.</given-names></name> <name><surname>Nico</surname> <given-names>P. S.</given-names></name> <name><surname>Pett-Ridge</surname> <given-names>J.</given-names></name> <name><surname>Kleber</surname> <given-names>M.</given-names></name></person-group> (<year>2015</year>). <article-title>Mineral protection of soil carbon counteracted by root exudates</article-title>. <source>Nat. Clim. Chang.</source> <volume>5</volume>, <fpage>588</fpage>&#x2013;<lpage>595</lpage>. doi: <pub-id pub-id-type="doi">10.1038/NCLIMATE2580</pub-id></citation></ref>
<ref id="ref29"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lal</surname> <given-names>R.</given-names></name></person-group> (<year>2020</year>). <article-title>Soil organic matter content and crop yield</article-title>. <source>J. Soil Water Conserv.</source> <volume>75</volume>, <fpage>27A</fpage>&#x2013;<lpage>32A</lpage>. doi: <pub-id pub-id-type="doi">10.2489/jswc.75.2.27A</pub-id> %J.</citation></ref>
<ref id="ref30"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lanoue</surname> <given-names>A.</given-names></name> <name><surname>Burlat</surname> <given-names>V.</given-names></name> <name><surname>Henkes</surname> <given-names>G. J.</given-names></name> <name><surname>Koch</surname> <given-names>I.</given-names></name> <name><surname>Schurr</surname> <given-names>U.</given-names></name> <name><surname>R&#x00F6;se</surname> <given-names>U. S. R.</given-names></name></person-group> (<year>2010</year>). <article-title>De novo biosynthesis of defense root exudates in response to fusarium attack in barley</article-title>. <source>New Phytol.</source> <volume>185</volume>, <fpage>577</fpage>&#x2013;<lpage>588</lpage>. doi: <pub-id pub-id-type="doi">10.1111/j.1469-8137.2009.03066.x</pub-id>, PMID: <pub-id pub-id-type="pmid">19878462</pub-id></citation></ref>
<ref id="ref9004"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Li</surname> <given-names>P. C.</given-names></name> <name><surname>Wang</surname> <given-names>S. L.</given-names></name> <name><surname>Qi</surname> <given-names>H.</given-names></name> <name><surname>Wang</surname> <given-names>Y.</given-names></name> <name><surname>Zhang</surname> <given-names>Q.</given-names></name> <name><surname>Feng</surname> <given-names>G. Y.</given-names></name> <etal/></person-group>. (<year>2019</year>). <article-title>Soil replacement combined with subsoiling improves cotton yields</article-title>. <source>J. Cotton Res.</source> <volume>2</volume>, <fpage>203</fpage>&#x2013;<lpage>218</lpage>. doi: <pub-id pub-id-type="doi">10.1186/s42397-019-0038-x</pub-id></citation></ref>
<ref id="ref31"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Li</surname> <given-names>X.</given-names></name> <name><surname>Thwe</surname> <given-names>A. A.</given-names></name> <name><surname>Park</surname> <given-names>N. I.</given-names></name> <name><surname>Suzuki</surname> <given-names>T.</given-names></name> <name><surname>Kim</surname> <given-names>S. J.</given-names></name> <name><surname>Park</surname> <given-names>S. U.</given-names></name></person-group> (<year>2012</year>). <article-title>Accumulation of Phenylpropanoids and correlated gene expression during the development of Tartary buckwheat sprouts</article-title>. <source>J. Agric. Food Chem.</source> <volume>60</volume>, <fpage>5629</fpage>&#x2013;<lpage>5635</lpage>. doi: <pub-id pub-id-type="doi">10.1021/jf301449a</pub-id>, PMID: <pub-id pub-id-type="pmid">22587625</pub-id></citation></ref>
<ref id="ref32"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Li</surname> <given-names>J.</given-names></name> <name><surname>Ye</surname> <given-names>X.</given-names></name> <name><surname>An</surname> <given-names>J.</given-names></name> <name><surname>Jin</surname> <given-names>X.</given-names></name> <name><surname>Fan</surname> <given-names>Q.</given-names></name> <name><surname>Zou</surname> <given-names>H.</given-names></name> <etal/></person-group>. (<year>2022</year>). <article-title>The more straw we deep-bury, the more soil TOC will be accumulated: when soil bacteria abundance keeps growing</article-title>. <source>J. Soils Sediments</source> <volume>22</volume>, <fpage>162</fpage>&#x2013;<lpage>171</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s11368-021-03068-w</pub-id></citation></ref>
<ref id="ref33"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Li</surname> <given-names>J.</given-names></name> <name><surname>Ye</surname> <given-names>X.</given-names></name> <name><surname>Zhang</surname> <given-names>Y.</given-names></name> <name><surname>Chen</surname> <given-names>J.</given-names></name> <name><surname>Yu</surname> <given-names>N.</given-names></name> <name><surname>Zou</surname> <given-names>H.</given-names></name></person-group> (<year>2021</year>). <article-title>Maize straw deep-burying promotes soil bacteria community abundance and improves soil fertility</article-title>. <source>J. Soil Sci. Plant Nutr.</source> <volume>21</volume>, <fpage>1397</fpage>&#x2013;<lpage>1407</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s42729-021-00448-6</pub-id></citation></ref>
<ref id="ref34"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Linlin</surname> <given-names>L. I.</given-names></name> <name><surname>Jianguo</surname> <given-names>L.</given-names></name> <name><surname>Peng</surname> <given-names>Y.</given-names></name> <name><surname>Rong</surname> <given-names>T.</given-names></name> <name><surname>Zhigui</surname> <given-names>B.</given-names></name> <name><surname>Wenliang</surname> <given-names>L.</given-names></name> <etal/></person-group>. (<year>2019</year>). <article-title>Allelopathic effects of different combination of phenolic acid allelochemicals on cotton seed germination and seedling growth</article-title>. <source>Sci. Rep.</source> <volume>38</volume>, <fpage>115</fpage>&#x2013;<lpage>159</lpage>. doi: <pub-id pub-id-type="doi">10.1038/s41598-021-83752-6</pub-id></citation></ref>
<ref id="ref35"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lozupone</surname> <given-names>C.</given-names></name> <name><surname>Lladser</surname> <given-names>M. E.</given-names></name> <name><surname>Knights</surname> <given-names>D.</given-names></name> <name><surname>Stombaugh</surname> <given-names>J.</given-names></name> <name><surname>Knight</surname> <given-names>R.</given-names></name></person-group> (<year>2011</year>). <article-title>Uni Frac: an effective distance metric for microbial community comparison</article-title>. <source>ISME J.</source> <volume>5</volume>, <fpage>169</fpage>&#x2013;<lpage>172</lpage>. doi: <pub-id pub-id-type="doi">10.1038/ismej.2010.133</pub-id>, PMID: <pub-id pub-id-type="pmid">20827291</pub-id></citation></ref>
<ref id="ref36"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lu</surname> <given-names>T.</given-names></name> <name><surname>Ke</surname> <given-names>M.</given-names></name> <name><surname>Lavoie</surname> <given-names>M.</given-names></name> <name><surname>Jin</surname> <given-names>Y.</given-names></name> <name><surname>Fan</surname> <given-names>X.</given-names></name> <name><surname>Zhang</surname> <given-names>Z.</given-names></name> <etal/></person-group>. (<year>2018</year>). <article-title>Rhizosphere microorganisms can influence the timing of plant flowering</article-title>. <source>Microbiome</source> <volume>6</volume>:<fpage>231</fpage>. doi: <pub-id pub-id-type="doi">10.1186/s40168-018-0615-0</pub-id>, PMID: <pub-id pub-id-type="pmid">30587246</pub-id></citation></ref>
<ref id="ref37"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Lyu</surname> <given-names>B. C.</given-names></name> <name><surname>Sun</surname> <given-names>H.</given-names></name> <name><surname>Qian</surname> <given-names>J. Q.</given-names></name> <name><surname>Liang</surname> <given-names>H.</given-names></name> <name><surname>Zhu</surname> <given-names>J. P.</given-names></name> <name><surname>Zhang</surname> <given-names>Q. S.</given-names></name> <etal/></person-group>. (<year>2024</year>). <article-title>Interaction between root exudates of medicinal plants and rhizosphere microorganisms and its application in ecological planting of Chinese medicinal materials</article-title>. <source>Chin. J. Chin. Mater. Med.</source> <volume>49</volume>, <fpage>2128</fpage>&#x2013;<lpage>2137</lpage>. doi: <pub-id pub-id-type="doi">10.19540/j.cnki.cjcmm.20240119.102</pub-id>, PMID: <pub-id pub-id-type="pmid">38812228</pub-id></citation></ref>
<ref id="ref38"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Malik</surname> <given-names>H. M.</given-names></name> <name><surname>Jawad</surname> <given-names>K. A.</given-names></name></person-group> (<year>2022</year>). <article-title>Potassium forms status in some Iraqi sedimentary soils and effect of cultivation on it</article-title>. <source>IOP Conf. Ser. Earth Environ. Sci.</source> <volume>1060</volume>:<fpage>012018</fpage>. doi: <pub-id pub-id-type="doi">10.1088/1755-1315/1060/1/012018</pub-id></citation></ref>
<ref id="ref39"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Maurer</surname> <given-names>D.</given-names></name> <name><surname>Malique</surname> <given-names>F.</given-names></name> <name><surname>Alfarraj</surname> <given-names>S.</given-names></name> <name><surname>Albasher</surname> <given-names>G.</given-names></name> <name><surname>Rennenberg</surname> <given-names>H.</given-names></name></person-group> (<year>2021</year>). <article-title>Interactive regulation of root exudation and rhizosphere denitrification by plant metabolite content and soil properties</article-title>. <source>Plant Soil</source> <volume>467</volume>, <fpage>107</fpage>&#x2013;<lpage>127</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s11104-021-05069-7</pub-id></citation></ref>
<ref id="ref40"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Meihong</surname> <given-names>S.</given-names></name> <name><surname>Yulian</surname> <given-names>G.</given-names></name> <name><surname>Caifeng</surname> <given-names>C.</given-names></name> <name><surname>Manjuan</surname> <given-names>A.</given-names></name> <name><surname>Bijian</surname> <given-names>Z.</given-names></name> <name><surname>Jie</surname> <given-names>X.</given-names></name> <etal/></person-group>. (<year>2019</year>). <article-title>Advances in studies on the effects of root exudates on adsorption and desorption of heavy metals in soil</article-title>. <source>J. Green Sci. Technol.</source> <volume>6</volume>, <fpage>81</fpage>&#x2013;<lpage>86</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.ecoenv.2024.117036</pub-id></citation></ref>
<ref id="ref41"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Morales</surname> <given-names>M. E.</given-names></name> <name><surname>Iocoli</surname> <given-names>G. A.</given-names></name> <name><surname>Allegrini</surname> <given-names>M.</given-names></name> <name><surname>Villamil</surname> <given-names>M. B.</given-names></name> <name><surname>Zabaloy</surname> <given-names>M. C.</given-names></name></person-group> (<year>2022</year>). <article-title>Response of root exudates and bacterial community to N fertilization and termination methods in <italic>Avena sativa</italic> L. as a winter cover crop model</article-title>. <source>Eur. J. Soil Biol.</source> <volume>114</volume>:<fpage>103453</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.ejsobi.2022.103453</pub-id></citation></ref>
<ref id="ref42"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Nannipieri</surname> <given-names>P.</given-names></name> <name><surname>Ascher</surname> <given-names>J.</given-names></name> <name><surname>Ceccherini</surname> <given-names>M. T.</given-names></name> <name><surname>Landi</surname> <given-names>L.</given-names></name> <name><surname>Valori</surname> <given-names>F.</given-names></name></person-group> (<year>2008</year>). <article-title>Effects of root exudates in microbial diversity and activity in rhizosphere soils</article-title>. <source>Soil. Biol</source> <volume>15</volume>, <fpage>339</fpage>&#x2013;<lpage>365</lpage>. doi: <pub-id pub-id-type="doi">10.1007/978-3-540-75575-3-14</pub-id></citation></ref>
<ref id="ref43"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Nicolini</surname> <given-names>F.</given-names></name> <name><surname>Topp</surname> <given-names>W.</given-names></name></person-group> (<year>2005</year>). <article-title>Soil properties in plantations of sessile oak (<italic>Quercus petraea</italic>) and red oak (<italic>Quercus rubra</italic>) in reclaimed lignite open-cast mines of the Rhineland</article-title>. <source>Geoderma</source> <volume>129</volume>, <fpage>65</fpage>&#x2013;<lpage>72</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.geoderma.2004.12.031</pub-id></citation></ref>
<ref id="ref44"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Parray</surname> <given-names>J. A.</given-names></name> <name><surname>Jan</surname> <given-names>S.</given-names></name> <name><surname>Kamili</surname> <given-names>A. N.</given-names></name> <name><surname>Qadri</surname> <given-names>R. A.</given-names></name> <name><surname>Egamberdieva</surname> <given-names>D.</given-names></name> <name><surname>Ahmad</surname> <given-names>P.</given-names></name></person-group> (<year>2016</year>). <article-title>Current perspectives on plant growth-promoting rhizobacteria</article-title>. <source>J. Plant Growth Regul.</source> <volume>35</volume>, <fpage>877</fpage>&#x2013;<lpage>902</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s00344-016-9583-4</pub-id></citation></ref>
<ref id="ref45"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Pausch</surname> <given-names>J.</given-names></name> <name><surname>Kuzyakov</surname> <given-names>Y.</given-names></name></person-group> (<year>2018</year>). <article-title>Carbon input by roots into the soil: quantification of rhizodeposition from root to ecosystem scale</article-title>. <source>Glob. Change Biol.</source> <volume>24</volume>, <fpage>1</fpage>&#x2013;<lpage>12</lpage>. doi: <pub-id pub-id-type="doi">10.1111/gcb.13850</pub-id>, PMID: <pub-id pub-id-type="pmid">28752603</pub-id></citation></ref>
<ref id="ref46"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Qu</surname> <given-names>Q.</given-names></name> <name><surname>Zhang</surname> <given-names>Z.</given-names></name> <name><surname>Peijnenburg</surname> <given-names>W. J. G. M.</given-names></name> <name><surname>Liu</surname> <given-names>W.</given-names></name> <name><surname>Lu</surname> <given-names>T.</given-names></name> <name><surname>Hu</surname> <given-names>B.</given-names></name> <etal/></person-group>. (<year>2020</year>). <article-title>Rhizosphere microbiome assembly and its ipact on plant growth</article-title>. <source>J. Agric. Food Chem.</source> <volume>68</volume>, <fpage>5024</fpage>&#x2013;<lpage>5038</lpage>. doi: <pub-id pub-id-type="doi">10.1021/acs.jafc.0c00073</pub-id></citation></ref>
<ref id="ref47"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Rahman</surname> <given-names>M. K. U.</given-names></name> <name><surname>Wang</surname> <given-names>X.</given-names></name> <name><surname>Gao</surname> <given-names>D.</given-names></name> <name><surname>Zhou</surname> <given-names>X.</given-names></name> <name><surname>Wu</surname> <given-names>F.</given-names></name></person-group> (<year>2021</year>). <article-title>Root exudates increase phosphorus availability in the tomato/potato onion intercropping system</article-title>. <source>Plant Soil</source> <volume>464</volume>, <fpage>45</fpage>&#x2013;<lpage>62</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s11104-021-04935-8</pub-id></citation></ref>
<ref id="ref48"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Robson</surname> <given-names>T.</given-names></name> <name><surname>Golos</surname> <given-names>P. J.</given-names></name> <name><surname>Stevens</surname> <given-names>J.</given-names></name> <name><surname>Reid</surname> <given-names>N.</given-names></name></person-group> (<year>2018</year>). <article-title>Enhancing tailings revegetation using shallow cover systems in arid environments: hydrogeochemical, nutritional, and ecophysiological constraints</article-title>. <source>Land Degrad. Dev.</source> <volume>29</volume>, <fpage>2785</fpage>&#x2013;<lpage>2796</lpage>. doi: <pub-id pub-id-type="doi">10.1002/ldr.2980</pub-id></citation></ref>
<ref id="ref49"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Rosier</surname> <given-names>A.</given-names></name> <name><surname>Medeiros</surname> <given-names>F. H. V.</given-names></name> <name><surname>Bais</surname> <given-names>H. P.</given-names></name></person-group> (<year>2018</year>). <article-title>Defining plant growth promoting rhizobacteria molecular and biochemical networks in beneficial plant-microbe interactions</article-title>. <source>Plant Soil</source> <volume>428</volume>, <fpage>35</fpage>&#x2013;<lpage>55</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s11104-018-3679-5</pub-id></citation></ref>
<ref id="ref9006"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Shen</surname> <given-names>Q. S.</given-names></name> <name><surname>Zhang</surname> <given-names>L.</given-names></name> <name><surname>Ismael Wang</surname> <given-names>Z. D.</given-names></name> <name><surname>Gao</surname> <given-names>Q.</given-names></name> <name><surname>Chen</surname> <given-names>S.</given-names></name></person-group> (<year>2019</year>). <source>J. Oceanol. Limnol.</source> <volume>37</volume>. doi: <pub-id pub-id-type="doi">10.1007/s00343-019-7351-6</pub-id></citation></ref>
<ref id="ref50"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Speirs</surname> <given-names>L. B. M.</given-names></name> <name><surname>Rice</surname> <given-names>D. T. F.</given-names></name> <name><surname>Petrovski</surname> <given-names>S.</given-names></name> <name><surname>Seviour</surname> <given-names>R. J.</given-names></name></person-group> (<year>2019</year>). <article-title>The phylogeny, biodiversity, and ecology of the Chloroflexi in activated sludge</article-title>. <source>Front. Microbiol.</source> <volume>10</volume>:<fpage>2015</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fmicb.2019.02015</pub-id>, PMID: <pub-id pub-id-type="pmid">31572309</pub-id></citation></ref>
<ref id="ref51"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Sun</surname> <given-names>L.</given-names></name> <name><surname>Ataka</surname> <given-names>M.</given-names></name> <name><surname>Kominami</surname> <given-names>Y.</given-names></name> <name><surname>Yoshimura</surname> <given-names>K.</given-names></name> <name><surname>Kitayama</surname> <given-names>K.</given-names></name></person-group> (<year>2021</year>). <article-title>A constant microbial C/N ratio mediates the microbial nitrogen mineralization induced by root exudation among four co-existing canopy species</article-title>. <source>Rhizospher-Neth</source> <volume>17</volume>:<fpage>100317</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.rhisph.2021.100317</pub-id></citation></ref>
<ref id="ref52"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Sun</surname> <given-names>W.</given-names></name> <name><surname>Xiao</surname> <given-names>E.</given-names></name> <name><surname>Krumins</surname> <given-names>V.</given-names></name> <name><surname>H&#x00E4;ggblom</surname> <given-names>M. M.</given-names></name> <name><surname>Dong</surname> <given-names>Y.</given-names></name> <name><surname>Pu</surname> <given-names>Z.</given-names></name> <etal/></person-group>. (<year>2018</year>). <article-title>Rhizosphere microbial response to multiple metal (loid) s in different contaminated arable soils indicates crop-specific metal-microbe interactions</article-title>. <source>Appl. Environ. Microbiol.</source> <volume>84</volume>, <fpage>e00701</fpage>&#x2013;<lpage>e00718</lpage>. doi: <pub-id pub-id-type="doi">10.1128/AEM.00701-18</pub-id>, PMID: <pub-id pub-id-type="pmid">30291123</pub-id></citation></ref>
<ref id="ref53"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Tittonell</surname> <given-names>P.</given-names></name> <name><surname>Corbeels</surname> <given-names>M.</given-names></name> <name><surname>Van Wijk</surname> <given-names>M. T.</given-names></name> <name><surname>Vanlauwe</surname> <given-names>B.</given-names></name> <name><surname>Giller</surname> <given-names>K. E.</given-names></name></person-group> (<year>2008</year>). <article-title>Combining organic and mineral fertilizers for integrated soil fertility management in smallholder farming systems of Kenya: explorations using the crop-soil model FIELD</article-title>. <source>Agron. J.</source> <volume>100</volume>, <fpage>1511</fpage>&#x2013;<lpage>1526</lpage>. doi: <pub-id pub-id-type="doi">10.2134/agronj2007.0355</pub-id></citation></ref>
<ref id="ref54"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Tong</surname> <given-names>Y.</given-names></name> <name><surname>Zheng</surname> <given-names>X.</given-names></name> <name><surname>Hu</surname> <given-names>Y.</given-names></name> <name><surname>Wu</surname> <given-names>J.</given-names></name> <name><surname>Liu</surname> <given-names>H.</given-names></name> <name><surname>Deng</surname> <given-names>Y.</given-names></name> <etal/></person-group>. (<year>2024</year>). <article-title>Root exudate-mediated plant-microbiome interactions determine plant health during disease infection</article-title>. <source>Agric. Ecosyst. Environ.</source> <volume>370</volume>:<fpage>109056</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.agee.2024.109056</pub-id></citation></ref>
<ref id="ref55"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Ulbrich</surname> <given-names>T. C.</given-names></name> <name><surname>Rivas-Ubach</surname> <given-names>A.</given-names></name> <name><surname>Tiemann</surname> <given-names>L. K.</given-names></name> <name><surname>Friesen</surname> <given-names>M. L.</given-names></name> <name><surname>Evans</surname> <given-names>S. E.</given-names></name></person-group> (<year>2022</year>). <article-title>Plant root exudates and rhizosphere bacterial communities shift with neighbor context</article-title>. <source>Soil Biol. Biochem.</source> <volume>172</volume>:<fpage>108753</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.soilbio.2022.108753</pub-id></citation></ref>
<ref id="ref56"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Vives-Peris</surname> <given-names>V.</given-names></name> <name><surname>de Ollas</surname> <given-names>C.</given-names></name> <name><surname>G&#x00F3;mez-Cadenas</surname> <given-names>A.</given-names></name> <name><surname>P&#x00E9;rez-Clemente</surname> <given-names>R. M.</given-names></name></person-group> (<year>2020</year>). <article-title>Root exudates: from plant to rhizosphere and beyond</article-title>. <source>Plant Cell Rep.</source> <volume>39</volume>, <fpage>3</fpage>&#x2013;<lpage>17</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s00299-019-02447-5</pub-id>, PMID: <pub-id pub-id-type="pmid">31346716</pub-id></citation></ref>
<ref id="ref57"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Viviane</surname> <given-names>C.</given-names></name> <name><surname>Cristina</surname> <given-names>R.</given-names></name> <name><surname>Francisco</surname> <given-names>D. A.</given-names></name> <etal/></person-group>. (<year>2021</year>). <article-title>Successive plant growth amplifies genotype-specific assembly of the tomato rhizosphere microbiome</article-title>. <source>Sci. Total Environ.</source> <volume>772</volume>:<fpage>144825</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.scitotenv.2020.144825</pub-id>, PMID: <pub-id pub-id-type="pmid">33581524</pub-id></citation></ref>
<ref id="ref9005"><citation citation-type="other"><person-group person-group-type="author"><name><surname>Wan</surname> <given-names>S.</given-names></name> <name><surname>Mou</surname> <given-names>X. J.</given-names></name> <name><surname>Liu</surname> <given-names>X. T.</given-names></name></person-group> (<year>2018</year>). <article-title>Effects of Reclamation on Soil Carbon and Nitrogen in Coastal Wetlands of Liaohe River Delta</article-title>, <source>China. Chin. Geogr. Sci.</source> <volume>28</volume>, <fpage>443</fpage>&#x2013;<lpage>455</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s11769-018-0961-7</pub-id></citation></ref>
<ref id="ref58"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Wang</surname> <given-names>Y.</given-names></name> <name><surname>Luo</surname> <given-names>D.</given-names></name> <name><surname>Xiong</surname> <given-names>Z.</given-names></name> <name><surname>Wang</surname> <given-names>Z.</given-names></name> <name><surname>Gao</surname> <given-names>M.</given-names></name></person-group> (<year>2023</year>). <article-title>Changes in rhizosphere phosphorus fractions and phosphate-mineralizing microbial populations in acid soil as influenced by organic acid exudation</article-title>. <source>Soil Tillage Res.</source> <volume>225</volume>:<fpage>105543</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.still.2022.105543</pub-id></citation></ref>
<ref id="ref59"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Wang</surname> <given-names>Q.</given-names></name> <name><surname>Sun</surname> <given-names>H.</given-names></name> <name><surname>Xu</surname> <given-names>C.</given-names></name> <name><surname>Ma</surname> <given-names>L.</given-names></name> <name><surname>Li</surname> <given-names>M.</given-names></name> <name><surname>Shao</surname> <given-names>C.</given-names></name> <etal/></person-group>. (<year>2019</year>). <article-title>Analysis of rhizosphere bacterial and fungal communities associated with rusty root disease of <italic>Panax ginseng</italic></article-title>. <source>Appl. Soil Ecol.</source> <volume>138</volume>, <fpage>245</fpage>&#x2013;<lpage>252</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.apsoil.2019.03.012</pub-id></citation></ref>
<ref id="ref60"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Wang</surname> <given-names>L.</given-names></name> <name><surname>Wang</surname> <given-names>F.</given-names></name> <name><surname>Wang</surname> <given-names>S.</given-names></name> <name><surname>Huang</surname> <given-names>Y.</given-names></name> <name><surname>Zhang</surname> <given-names>Z.</given-names></name> <name><surname>Bai</surname> <given-names>Z.</given-names></name> <etal/></person-group>. (<year>2021</year>). <article-title>Analysis of differences in chemical properties of reconstructed soil under different proportions of topsoil substitute materials</article-title>. <source>Environ. Pollut.</source> <volume>28</volume>, <fpage>31230</fpage>&#x2013;<lpage>31245</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s11356-021-12803-2</pub-id>, PMID: <pub-id pub-id-type="pmid">33599929</pub-id></citation></ref>
<ref id="ref61"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Wang</surname> <given-names>Q.</given-names></name> <name><surname>Zhao</surname> <given-names>Y.</given-names></name> <name><surname>Xiao</surname> <given-names>W.</given-names></name></person-group> (<year>2022</year>). <article-title>Assessing potential spontaneous combustion of coal gangue dumps after reclamation by simulating alfalfa heat stress based on the spectral features of chlorophyll fluorescence parameters</article-title>. <source>REMOTE. SENS-BASEL.</source> <volume>14</volume>, <fpage>59</fpage>&#x2013;<lpage>74</lpage>. doi: <pub-id pub-id-type="doi">10.3390/rs14235974</pub-id></citation></ref>
<ref id="ref9007"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Wang</surname> <given-names>X. Y.</given-names></name> <name><surname>Jiang</surname> <given-names>J. P.</given-names></name> <name><surname>Hu</surname> <given-names>J. H.</given-names></name></person-group> (<year>2024</year>). <article-title>Spatiotemporal Shifts in the Richness and Composition of Mountain Frogs in a Global Biodiversity Hotspot in Southwest China</article-title>. <source>Asian Herpetol. Res.</source> <volume>15</volume>, <fpage>241</fpage>&#x2013;<lpage>250</lpage>. doi: <pub-id pub-id-type="doi">10.3724/ahr.2095-0357.2024.0024</pub-id></citation></ref>
<ref id="ref62"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Wen</surname> <given-names>Y. C.</given-names></name> <name><surname>Li</surname> <given-names>H. Y.</given-names></name> <name><surname>Lin</surname> <given-names>Z. A.</given-names></name> <name><surname>Zhao</surname> <given-names>B. Q.</given-names></name> <name><surname>Li</surname> <given-names>Y. Q.</given-names></name></person-group> (<year>2020</year>). <article-title>Long-term fertilization alters soil properties and fungal community composition in fluvo-aquic soil of the North China plain</article-title>. <source>Sci. Rep.</source> <volume>10</volume>:<fpage>7198</fpage>. doi: <pub-id pub-id-type="doi">10.1038/s41598-020-64227-6</pub-id>, PMID: <pub-id pub-id-type="pmid">32350351</pub-id></citation></ref>
<ref id="ref63"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Wilson-Kokes</surname> <given-names>L.</given-names></name> <name><surname>Emerson</surname> <given-names>P.</given-names></name> <name><surname>DeLong</surname> <given-names>C.</given-names></name> <name><surname>Thomas</surname> <given-names>C.</given-names></name> <name><surname>Skousen</surname> <given-names>J.</given-names></name></person-group> (<year>2013</year>). <article-title>Hardwood tree growth after eight years on brown and gray mine soils in West Virginia</article-title>. <source>J. Environ. Qual.</source> <volume>42</volume>, <fpage>1353</fpage>&#x2013;<lpage>1362</lpage>. doi: <pub-id pub-id-type="doi">10.2134/jeq2013.04.0113</pub-id>, PMID: <pub-id pub-id-type="pmid">24216413</pub-id></citation></ref>
<ref id="ref64"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Wu</surname> <given-names>Z.</given-names></name> <name><surname>Cui</surname> <given-names>F.</given-names></name> <name><surname>Nie</surname> <given-names>J.</given-names></name></person-group> (<year>2023</year>). <article-title>Relationship between soil water content and vegetation distribution in a small area before and after coal seam mining: a case study of coal mining subsidence area in Northwest China</article-title>. <source>Environ. Earth Sci.</source> <volume>1</volume>, <fpage>36</fpage>&#x2013;<lpage>41</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s12665-023-10791-9</pub-id></citation></ref>
<ref id="ref65"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Xie</surname> <given-names>L.</given-names></name> <name><surname>Li</surname> <given-names>W.</given-names></name> <name><surname>Pang</surname> <given-names>X.</given-names></name> <name><surname>Liu</surname> <given-names>Q.</given-names></name> <name><surname>Yin</surname> <given-names>C.</given-names></name></person-group> (<year>2022</year>). <article-title>Soil properties and root traits are important factors driving rhizosphere soil bacterial and fungal community variations in alpine <italic>Rhododendron nitidulum</italic> shrub ecosystems along an altitudinal gradient</article-title>. <source>Sci. Total Environ.</source> <volume>864</volume>:<fpage>161048</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.scitotenv.2022.161048</pub-id></citation></ref>
<ref id="ref66"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Xie</surname> <given-names>W. Y.</given-names></name> <name><surname>Yuan</surname> <given-names>S. T.</given-names></name> <name><surname>Xu</surname> <given-names>M. G.</given-names></name> <name><surname>Yang</surname> <given-names>X. P.</given-names></name> <name><surname>Shen</surname> <given-names>Q. R.</given-names></name> <name><surname>Zhang</surname> <given-names>W. W.</given-names></name> <etal/></person-group>. (<year>2018</year>). <article-title>Long-term effects of manure and chemical fertilizers on soil antibiotic resistome</article-title>. <source>Soil Biol. Biochem.</source> <volume>122</volume>, <fpage>111</fpage>&#x2013;<lpage>119</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.soilbio.2018.04.009</pub-id></citation></ref>
<ref id="ref67"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Xu</surname> <given-names>N.</given-names></name> <name><surname>Tan</surname> <given-names>G.</given-names></name> <name><surname>Wang</surname> <given-names>H.</given-names></name> <name><surname>Gai</surname> <given-names>X.</given-names></name></person-group> (<year>2016</year>). <article-title>Effect of biochar additions to soil on nitrogen leaching, microbial biomass and bacterial community structure</article-title>. <source>Eur. J. Soil Biol.</source> <volume>74</volume>, <fpage>1</fpage>&#x2013;<lpage>8</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.ejsobi.2016.02.004</pub-id></citation></ref>
<ref id="ref9008"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Yang</surname> <given-names>X. M.</given-names></name> <name><surname>Feng</surname> <given-names>Q.</given-names></name> <name><surname>Zhu</surname> <given-names>M.</given-names></name></person-group> (<year>2024</year>). <article-title>Vegetation characteristics and soil properties of artificially remediated grasslands: The case study of the Shimenhe mining area in Qilian Mountains, northwest China</article-title>. <source>Res Cold Arid Reg.</source> <volume>16</volume>, <fpage>190</fpage>&#x2013;<lpage>200</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.rcar.2024.09.001</pub-id></citation></ref>
<ref id="ref68"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Zeng</surname> <given-names>F.</given-names></name> <name><surname>Chen</surname> <given-names>S.</given-names></name> <name><surname>Miao</surname> <given-names>Y.</given-names></name> <name><surname>Wu</surname> <given-names>F.</given-names></name> <name><surname>Zhang</surname> <given-names>G.</given-names></name></person-group> (<year>2008</year>). <article-title>Changes of organic acid exudation and rhizosphere pH in rice plants under chromium stress</article-title>. <source>Environ. Pollut.</source> <volume>155</volume>, <fpage>284</fpage>&#x2013;<lpage>289</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.envpol.2007.11.019</pub-id>, PMID: <pub-id pub-id-type="pmid">18162271</pub-id></citation></ref>
<ref id="ref69"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Zhang</surname> <given-names>S.</given-names></name> <name><surname>Li</surname> <given-names>X.</given-names></name> <name><surname>Chen</surname> <given-names>K.</given-names></name> <name><surname>Shi</surname> <given-names>J.</given-names></name> <name><surname>Wang</surname> <given-names>Y.</given-names></name> <name><surname>Luo</surname> <given-names>P.</given-names></name> <etal/></person-group>. (<year>2022</year>). <article-title>Long-term fertilization altered microbial community structure in an aeolian sandy soil in Northeast China</article-title>. <source>Front. Microbiol.</source> <volume>13</volume>:<fpage>979759</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fmicb.2022.979759</pub-id>, PMID: <pub-id pub-id-type="pmid">36160213</pub-id></citation></ref>
<ref id="ref70"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Zhao</surname> <given-names>D.</given-names></name> <name><surname>Hou</surname> <given-names>H.</given-names></name> <name><surname>Liu</surname> <given-names>H.</given-names></name></person-group> (<year>2023</year>). <article-title>Microbial community structure and predictive functional analysis in reclaimed soil with different vegetation types: the example of the Xiaoyi mine waste dump in Shanxi</article-title>. <source>Land</source> <volume>12</volume>, <fpage>456</fpage>&#x2013;<lpage>459</lpage>. doi: <pub-id pub-id-type="doi">10.3390/land12020456</pub-id></citation></ref>
<ref id="ref71"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Zhao</surname> <given-names>M.</given-names></name> <name><surname>Zhao</surname> <given-names>J.</given-names></name> <name><surname>Yuan</surname> <given-names>J.</given-names></name> <name><surname>Hale</surname> <given-names>L.</given-names></name> <name><surname>Wen</surname> <given-names>T.</given-names></name> <name><surname>Huang</surname> <given-names>Q.</given-names></name> <etal/></person-group>. (<year>2025</year>). <article-title>Root exudates drive soil-microbe-nutrient feedbacks in response to plant growth</article-title>. <source>Plant Cell Environ.</source> <volume>44</volume>, <fpage>613</fpage>&#x2013;<lpage>628</lpage>. doi: <pub-id pub-id-type="doi">10.1111/pce.13928</pub-id></citation></ref>
<ref id="ref72"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Zhou</surname> <given-names>X.</given-names></name> <name><surname>Yu</surname> <given-names>G.</given-names></name> <name><surname>Wu</surname> <given-names>F.</given-names></name></person-group> (<year>2012</year>). <article-title>Soil phenolics in a continuously mono-cropped cucumber (<italic>Cucumis sativus</italic> L.) system and their effects on cucumber seedling growth and soil microbial communities</article-title>. <source>Eur. J. Soil Sci.</source> <volume>63</volume>, <fpage>332</fpage>&#x2013;<lpage>340</lpage>. doi: <pub-id pub-id-type="doi">10.1111/j.1365-2389.2012.01442.x</pub-id></citation></ref>
<ref id="ref73"><citation citation-type="journal"><person-group person-group-type="author"><name><surname>Zhou</surname> <given-names>X.</given-names></name> <name><surname>Zhang</surname> <given-names>J.</given-names></name> <name><surname>Pan</surname> <given-names>D.</given-names></name> <name><surname>Ge</surname> <given-names>X.</given-names></name> <name><surname>Jin</surname> <given-names>X.</given-names></name> <name><surname>Chen</surname> <given-names>S.</given-names></name> <etal/></person-group>. (<year>2018</year>). <article-title>P-coumaric can alter the composition of cucumber rhizosphere microbial communities and induce negative plant-microbial interactions</article-title>. <source>Biol. Fertil. Soils</source> <volume>54</volume>, <fpage>363</fpage>&#x2013;<lpage>372</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s00374-018-1265-x</pub-id></citation></ref>
</ref-list>
</back>
</article>