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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Microbiol.</journal-id>
<journal-title-group>
<journal-title>Frontiers in Microbiology</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Microbiol.</abbrev-journal-title>
</journal-title-group>
<issn pub-type="epub">1664-302X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fmicb.2026.1771649</article-id>
<article-version article-version-type="Version of Record" vocab="NISO-RP-8-2008"/>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Review</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>Microbial biocontrol agents and the rhizosphere microbiome: integrating ecological function and climate resilience in sustainable agriculture</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Iqbal</surname>
<given-names>Mudassir</given-names>
</name>
<xref ref-type="aff" rid="aff1"/>
<xref ref-type="corresp" rid="c001"><sup>&#x002A;</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/2025183"/>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="conceptualization" vocab-term-identifier="https://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Data curation" vocab-term-identifier="https://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Funding acquisition" vocab-term-identifier="https://credit.niso.org/contributor-roles/funding-acquisition/">Funding acquisition</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="investigation" vocab-term-identifier="https://credit.niso.org/contributor-roles/investigation/">Investigation</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="methodology" vocab-term-identifier="https://credit.niso.org/contributor-roles/methodology/">Methodology</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Project administration" vocab-term-identifier="https://credit.niso.org/contributor-roles/project-administration/">Project administration</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="resources" vocab-term-identifier="https://credit.niso.org/contributor-roles/resources/">Resources</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; original draft" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-original-draft/">Writing &#x2013; original draft</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; review &#x0026; editing" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-review-editing/">Writing &#x2013; review &#x0026; editing</role>
</contrib>
</contrib-group>
<aff id="aff1"><institution>Department of Plant Protection Biology, Swedish University of Agricultural Sciences</institution>, <city>Lomma</city>, <country country="se">Sweden</country></aff>
<author-notes>
<corresp id="c001"><label>&#x002A;</label>Correspondence: Mudassir Iqbal, <email xlink:href="mailto:mudassir.iqbal@slu.se">mudassir.iqbal@slu.se</email></corresp>
</author-notes>
<pub-date publication-format="electronic" date-type="pub" iso-8601-date="2026-02-26">
<day>26</day>
<month>02</month>
<year>2026</year>
</pub-date>
<pub-date publication-format="electronic" date-type="collection">
<year>2026</year>
</pub-date>
<volume>17</volume>
<elocation-id>1771649</elocation-id>
<history>
<date date-type="received">
<day>19</day>
<month>12</month>
<year>2025</year>
</date>
<date date-type="rev-recd">
<day>28</day>
<month>01</month>
<year>2026</year>
</date>
<date date-type="accepted">
<day>16</day>
<month>02</month>
<year>2026</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x00A9; 2026 Iqbal.</copyright-statement>
<copyright-year>2026</copyright-year>
<copyright-holder>Iqbal</copyright-holder>
<license>
<ali:license_ref start_date="2026-02-26">https://creativecommons.org/licenses/by/4.0/</ali:license_ref>
<license-p>This is an open-access article distributed under the terms of the <ext-link ext-link-type="uri" xlink:href="https://creativecommons.org/licenses/by/4.0/">Creative Commons Attribution License (CC BY)</ext-link>. The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</license-p>
</license>
</permissions>
<abstract>
<p>The growing challenges of food insecurity, soil degradation, and climate-induced stresses are exposing the limitations of chemically intensive crop protection systems. In this context, the rhizosphere microbiome, comprising complex microbial networks that regulate plant growth, nutrient acquisition, and immune responses, has emerged as a promising focus for more sustainable agricultural practices. Microbial biocontrol agents (BCAs) are increasingly recognized not only for their pathogen-suppressive properties but also for their potential to modulate rhizosphere microbial communities and contribute to plant tolerance to abiotic stressors. This review synthesizes recent advances in understanding the ecological and mechanistic interplay between BCAs and the rhizosphere microbiome, highlighting how microbial inoculants can influence community assembly, functional processes, and microbiome resilience under biotic and abiotic stress conditions. Drawing on molecular and ecological evidence, the synthesis integrates current knowledge of BCA-mediated regulation of plant defense signalling, nutrient cycling, and stress-associated responses. Key knowledge gaps related to inoculant persistence, ecological compatibility, and microbiome-level trade-offs that limit field-scale effectiveness are also identified. To address these challenges, a microbiome-informed conceptual framework is proposed, emphasizing precision-designed synthetic microbial communities (SynComs), trait-based screening, host&#x2013;microbiome co-optimization, and integration of BCAs into resilient Integrated Pest Management (IPM) strategies. In summary, this review provides a systems-level perspective on how rhizosphere microbiome dynamics can be leveraged to support sustainable climate-smart crop production.</p>
</abstract>
<kwd-group>
<kwd>biocontrol agents</kwd>
<kwd>integrated pest management</kwd>
<kwd>microbial consortia</kwd>
<kwd>plant&#x2013;microbe interactions</kwd>
<kwd>rhizosphere microbiome</kwd>
<kwd>soil health</kwd>
</kwd-group>
<funding-group>
<funding-statement>The author(s) declared that financial support was received for this work and/or its publication. This work was supported by The Crafoord Foundation (grant no. 20240795), Novo Nordisk Foundation (grant no. NNF25OC0106993), the Royal Swedish Academy of Agriculture and Forestry (KSLA, grant no. CF2024-0013), the Martha och Dagny Larssons (SLU-LTV, 2025), and the Ekhaga Foundation (grant no. 2025-87).</funding-statement>
</funding-group>
<counts>
<fig-count count="1"/>
<table-count count="4"/>
<equation-count count="0"/>
<ref-count count="156"/>
<page-count count="15"/>
<word-count count="13375"/>
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<custom-meta-group>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Microbe and Virus Interactions with Plants</meta-value>
</custom-meta>
</custom-meta-group>
</article-meta>
</front>
<body>
<sec sec-type="intro" id="sec1">
<label>1</label>
<title>Introduction</title>
<p>Agricultural systems worldwide are facing challenges in ensuring food security for a rapidly growing population, which is expected to exceed 9.7 billion by 2050 (<xref ref-type="bibr" rid="ref131">Suman et al., 2022</xref>; <xref ref-type="bibr" rid="ref141">United Nations, 2019</xref>). To meet this demand, an estimated 70% increase in food production will be required (<xref ref-type="bibr" rid="ref105">Poveda, 2021</xref>). However, agriculture must rely on limited cultivable land that is increasingly threatened by climate change, land degradation associated with declining soil function, and unsustainable farming practices. Climate-induced abiotic stressors such as drought, erratic rainfall, rising temperatures, and soil salinization are already disrupting plant physiological processes and reducing yields (<xref ref-type="bibr" rid="ref97">Park et al., 2023</xref>; <xref ref-type="bibr" rid="ref151">Zandalinas et al., 2022</xref>). At the same time, shifting environmental conditions are reshaping plant&#x2013;pathogen interactions, accelerating the spread, persistence, and virulence of destructive plant pathogens including bacteria, fungi, oomycetes, viruses, and nematodes (<xref ref-type="bibr" rid="ref126">Singh et al., 2023</xref>). Collectively, these biotic stressors contribute to an estimated $220 billion in annual crop losses (<xref ref-type="bibr" rid="ref114">Ristaino et al., 2021</xref>; <xref ref-type="bibr" rid="ref116">Rohr et al., 2019</xref>; <xref ref-type="bibr" rid="ref142">Van Dijk et al., 2021</xref>) that is further exacerbated by post-harvest deterioration (<xref ref-type="bibr" rid="ref128">Srivastava, 2019</xref>), posing a direct threat to global food security.</p>
<p>While current management relies heavily on synthetic chemical inputs, these conventional approaches are increasingly proving inadequate under escalating biotic and abiotic pressures. These limitations are not only due to declining efficacy and the rise of resistant pathogen strains (<xref ref-type="bibr" rid="ref33">Fisher et al., 2018</xref>; <xref ref-type="bibr" rid="ref74">Lucas et al., 2015</xref>), but also to their unintended consequences for environmental health (<xref ref-type="bibr" rid="ref98">Pathak et al., 2022</xref>), biodiversity (<xref ref-type="bibr" rid="ref121">S&#x00E1;nchez-Bayo and Wyckhuys, 2019</xref>), and human well-being (<xref ref-type="bibr" rid="ref7">Asghar et al., 2016</xref>). The overreliance on synthetic pesticides and fertilizers has been linked to soil microbiome disruption, contamination of water bodies, bioaccumulation in food chains, and negative impacts on pollinators and beneficial organisms (<xref ref-type="bibr" rid="ref28">Devi et al., 2018</xref>; <xref ref-type="bibr" rid="ref37">Goulson et al., 2015</xref>). Additionally, these practices raise serious concerns about food safety and long-term sustainability, necessitating the urgent development of ecologically sound alternatives.</p>
<p>As a result, growing attention has turned to the rhizosphere microbiome, a dense and dynamic microbial consortium at the soil&#x2013;root interface that functions as a fundamental determinant of plant health, stress tolerance, and disease resistance (<xref ref-type="bibr" rid="ref102">Philippot et al., 2013</xref>). These microbial communities reinforce a wide array of physiological and ecological functions, including nutrient cycling, phytohormone signalling, modulation of root architecture, and immune priming, while simultaneously acting as a biological barrier against soilborne pathogens (<xref ref-type="bibr" rid="ref13">Busby et al., 2017</xref>; <xref ref-type="bibr" rid="ref97">Park et al., 2023</xref>). The rhizosphere microbiome&#x2019;s sensitivity to environmental cues and high functional plasticity reflect its function as a buffer against climatic stressors and mark it out as a promising target for agroecological innovation (<xref ref-type="bibr" rid="ref139">Trivedi et al., 2022</xref>). Microbiome manipulation and targeted microbiome modulation are thus increasingly recognized as pivotal strategies for advancing climate-smart and resilient agricultural systems while avoiding the limitations of chemical-intensive practices (<xref ref-type="bibr" rid="ref42">Han and Yoshikuni, 2022</xref>; <xref ref-type="bibr" rid="ref59">Kaur et al., 2025</xref>). However, deeper knowledge is required to enable predictable, ecologically compatible, and field-relevant intentional microbiome manipulation.</p>
<p>While the rhizosphere microbiome&#x2019;s central role in plant health and stress resilience has been recognized, the availability of practical and scalable tools for its manipulation is limited. Microbial biological control agents (BCAs) may represent one such tool. Several studies have examined the ability of microbial BCAs to suppress plant pathogens (<xref ref-type="bibr" rid="ref52">Iqbal et al., 2021</xref>, <xref ref-type="bibr" rid="ref50">2023</xref>; <xref ref-type="bibr" rid="ref136">Thangaraj et al., 2025</xref>) and the native microbiome&#x2019;s impact on crop performance (<xref ref-type="bibr" rid="ref13">Busby et al., 2017</xref>; <xref ref-type="bibr" rid="ref14">Chandel et al., 2019</xref>; <xref ref-type="bibr" rid="ref102">Philippot et al., 2013</xref>). Conversely, few have explored how BCAs interact with, restructure, or integrate into rhizosphere microbial communities to enhance plant resilience under combined biotic and abiotic stresses. Understanding of these issue is particularly important in the context of climate-resilient agriculture and integrated pest management (IPM) systems, where sustainability depends not only on effectiveness but also on ecological compatibility. In this context, ecological compatibility refers to the capacity of introduced BCAs to establish, persist, and function within native rhizosphere communities while maintaining beneficial microbial interactions and essential ecosystem processes (<xref ref-type="bibr" rid="ref87">Muhammad et al., 2024</xref>; <xref ref-type="bibr" rid="ref129">Stenberg, 2017</xref>; <xref ref-type="bibr" rid="ref146">Whipps, 2001</xref>). Despite its importance, the persistence, colonization, and functional influence of BCAs within native microbiomes remain poorly explored across crop systems and environmental gradients.</p>
<p>In this review, I synthesize current understanding of the ecological interactions between microbial BCAs and the rhizosphere microbiome. Specifically, I discuss the mechanisms through which BCAs influence microbial community assembly and function, highlight case studies illustrating crop-specific and context-dependent outcomes, and assess the implications for sustainable disease management and climate adaptation. Finally, I propose a conceptual framework for microbiome-informed biocontrol strategies (<xref ref-type="fig" rid="fig1">Figure 1</xref>) that align with the principles of IPM and climate-smart agriculture, integrating near-term deployable approaches with longer-term innovation pathways, and advancing a systems-level approach to enhancing crop resilience and sustainable productivity.</p>
<fig position="float" id="fig1">
<label>Figure 1</label>
<caption>
<p>Conceptual framework for microbiome-informed biological control in climate-smart agriculture. The framework illustrates an iterative, systems-level approach integrating consortium-level microbial design, trait-based screening and optimization, microbiome-informed integrated pest management, and host&#x2013;microbiome co-optimization. By explicitly incorporating microbial functional traits, ecological compatibility with resident microbiomes, and validation across environmental gradients, the framework extends conventional IPM strategies toward context-dependent, resilient disease management under climate stress.</p>
</caption>
<graphic xlink:href="fmicb-17-1771649-g001.tif" mimetype="image" mime-subtype="tiff">
<alt-text content-type="machine-generated">Cycle diagram illustrating microbiome-based precision agriculture strategies, including microbial design and SynComs, omics-based screening and synthetic biology, microbiome-informed IPM and precision agriculture, and host genetics and holobiont optimization, connected by arrows showing process flow.</alt-text>
</graphic>
</fig>
</sec>
<sec id="sec2">
<label>2</label>
<title>The rhizosphere microbiome: composition, function, and environmental sensitivity</title>
<sec id="sec3">
<label>2.1</label>
<title>Rhizosphere microbiome</title>
<p>The rhizosphere microbiome is structured by complex, dynamic interactions between plant genotype, soil characteristics, and environmental conditions. It plays an important role in determining plant health, nutrient cycling, and overall agroecosystem functionality (<xref ref-type="bibr" rid="ref13">Busby et al., 2017</xref>; <xref ref-type="bibr" rid="ref78">Massart et al., 2015</xref>). While soil is the primary reservoir of microbial inocula, its physicochemical properties such as pH, texture, organic matter content, and water-holding capacity strongly influence the baseline microbial community available for root colonization (<xref ref-type="bibr" rid="ref12">Berg and Smalla, 2009</xref>; <xref ref-type="bibr" rid="ref11">Berg et al., 2013</xref>). In the rhizosphere, this soil-derived pool is further filtered by plant-specific traits including root architecture, exudate composition, and immune signalling pathways. Experimental evidence demonstrates that even within identical soil types, different plant genotypes can assemble distinct microbial consortia, highlighting the host&#x2019;s regulatory role in microbiome assembly (<xref ref-type="bibr" rid="ref48">Ikeda et al., 2011</xref>; <xref ref-type="bibr" rid="ref97">Park et al., 2023</xref>).</p>
<p>In addition to inherent plant&#x2013;soil interactions, anthropogenic factors such as fertilization regimes, pesticides, and cropping patterns significantly shape rhizosphere microbiome structure and function. Fertilizers, particularly those applied in excess, have been linked to reduced microbial diversity and functional redundancy, potentially impairing key processes like nutrient cycling and pathogen suppression (<xref ref-type="bibr" rid="ref54">Jacobsen and Hjelms&#x00F8;, 2014</xref>). Conversely, practices such as crop rotation and organic amendments tend to promote microbial resilience, enhance beneficial taxa, and suppress soilborne pathogens (<xref ref-type="bibr" rid="ref46">Hilton et al., 2013</xref>). These findings challenge the concept of the rhizosphere as a passive microbial habitat and instead highlight its dynamic responsiveness to agronomic management. Within the broader framework of climate-smart agriculture, the rhizosphere represents not only a site of biological activity but also a manipulable interface through which plant resilience, productivity, and sustainability can be enhanced.</p>
</sec>
<sec id="sec4">
<label>2.2</label>
<title>Composition and key microbial groups</title>
<p>The rhizosphere microbiome is a phylogenetically diverse and functionally heterogeneous assemblage of microorganisms including bacteria, fungi, archaea, protists, and viruses that collectively shape plant&#x2013;soil interactions (<xref ref-type="bibr" rid="ref20">Chukwuneme and Babalola, 2025</xref>). These groups have complementary ecological functions that influence nutrient cycling, plant defense, and stress responses.</p>
<p>Bacteria are the most abundant and metabolically versatile members of the rhizosphere microbiome. Dominant phyla such as Proteobacteria, Actinobacteria, Firmicutes, and Bacteroidetes contribute to key functions including nutrient cycling, siderophore production, phytohormone modulation, and pathogen suppression (<xref ref-type="bibr" rid="ref31">Fierer, 2017</xref>; <xref ref-type="bibr" rid="ref66">Kour et al., 2023</xref>). Bacterial community composition is highly dynamic and shaped by host genotype, soil properties, and environmental conditions, with stress factors often favoring specific functional groups, such as Proteobacteria under nitrogen limitation and Actinobacteria under drought (<xref ref-type="bibr" rid="ref89">Negre Rodr&#x00ED;guez et al., 2025</xref>; <xref ref-type="bibr" rid="ref132">Sun et al., 2024</xref>). Within this context, keystone members of the rhizosphere microbiome include both functionally distinct groups and taxonomically defined but functionally versatile genera. For example, diazotrophs act as keystone functional groups through their contribution to biological nitrogen fixation (<xref ref-type="bibr" rid="ref49">Imran et al., 2021</xref>), whereas <italic>Pseudomonas</italic> spp. represent keystone genera that influence community structure and plant performance via pathogen suppression, siderophore production, and induced systemic resistance (<xref ref-type="bibr" rid="ref79">Mazzola, 2002</xref>; <xref ref-type="bibr" rid="ref139">Trivedi et al., 2022</xref>).</p>
<p>Fungi are another major functional component of the rhizosphere microbiome. Filamentous fungi from Ascomycota and Basidiomycota contribute to organic matter decomposition, phosphorus solubilization, and pathogen suppression through mycoparasitism and secondary metabolite production (<xref ref-type="bibr" rid="ref15">Chapelle et al., 2016</xref>; <xref ref-type="bibr" rid="ref112">Rigobelo, 2025</xref>). Arbuscular mycorrhizal fungi (AMF; Glomeromycota) form symbiotic associations with plant roots, enhancing nutrient acquisition, water-use efficiency, and tolerance of abiotic stresses such as drought and salinity while also influencing soil structure and host immune responses (<xref ref-type="bibr" rid="ref91">Nie et al., 2024</xref>; <xref ref-type="bibr" rid="ref144">Wahab et al., 2023</xref>). Fungal community composition is strongly shaped by plant genotype, soil pH, and nutrient availability.</p>
<p>Archaea, although generally less abundant than bacteria and fungi, play important roles in rhizosphere biogeochemical cycling, particularly in ammonia oxidation and anaerobic methanogenesis (<xref ref-type="bibr" rid="ref94">Offre et al., 2013</xref>; <xref ref-type="bibr" rid="ref107">Prosser, 2012</xref>). These processes are especially relevant in nutrient-poor, saline, or anoxic soils, where archaeal activity complements bacterial metabolism. Protists and viruses are key regulators of rhizosphere microbial dynamics. Protists influence microbial turnover through selective bacterivory, thereby affecting nutrient mineralization and community succession (<xref ref-type="bibr" rid="ref73">Liao et al., 2024</xref>). Bacteriophages shape bacterial populations through lytic and lysogenic interactions that regulate population dynamics and mediate horizontal gene transfer, thereby contributing to microbial adaptation and functional stability under changing environmental conditions (<xref ref-type="bibr" rid="ref18">Chevallereau et al., 2022</xref>). Collectively, interactions among these microbial guilds underpin the ecological functioning and resilience of the rhizosphere. <xref ref-type="table" rid="tab1">Table 1</xref> summarizes the major rhizosphere microbial groups, showing representative taxa from each and their dominant functional roles.</p>
<table-wrap position="float" id="tab1">
<label>Table 1</label>
<caption>
<p>Major microbial groups inhabiting the rhizosphere, representative taxa, and their dominant functional roles relevant to plant health, disease regulation, and stress resilience.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="top">Microbial groups</th>
<th align="left" valign="top">Representative taxa (examples)</th>
<th align="left" valign="top">Dominant functional roles</th>
<th align="left" valign="top">Relevance to plant health and stress</th>
<th align="left" valign="top">References</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="top">Beneficial bacteria</td>
<td align="left" valign="top"><italic>Pseudomonas, Bacillus, Azospirillum</italic></td>
<td align="left" valign="top">Nutrient solubilization, phytohormone modulation, ISR induction, antibiosis</td>
<td align="left" valign="top">Enhanced nutrient uptake, disease suppression, drought tolerance</td>
<td align="left" valign="top"><xref ref-type="bibr" rid="ref31">Fierer (2017)</xref>, <xref ref-type="bibr" rid="ref79">Mazzola (2002)</xref></td>
</tr>
<tr>
<td align="left" valign="top">Beneficial fungi</td>
<td align="left" valign="top"><italic>Trichoderma</italic>, AMF (Glomeromycota)</td>
<td align="left" valign="top">Mycoparasitism, nutrient acquisition, ISR, improved water uptake</td>
<td align="left" valign="top">Pathogen suppression, abiotic stress tolerance</td>
<td align="left" valign="top"><xref ref-type="bibr" rid="ref15">Chapelle et al. (2016)</xref>, <xref ref-type="bibr" rid="ref91">Nie et al. (2024)</xref></td>
</tr>
<tr>
<td align="left" valign="top">Plant pathogens</td>
<td align="left" valign="top"><italic>Fusarium, Ralstonia, Rhizoctonia</italic></td>
<td align="left" valign="top">Host colonization, microbiome disruption</td>
<td align="left" valign="top">Disease development, microbiome dysbiosis</td>
<td align="left" valign="top"><xref ref-type="bibr" rid="ref10">Berendsen et al. (2012)</xref>, <xref ref-type="bibr" rid="ref27">Deng et al. (2021)</xref>, <xref ref-type="bibr" rid="ref130">Su et al. (2020)</xref></td>
</tr>
<tr>
<td align="left" valign="top">Archaea</td>
<td align="left" valign="top">Ammonia-oxidizing archaea</td>
<td align="left" valign="top">Nitrogen cycling, stress-associated nutrient turnover</td>
<td align="left" valign="top">Nutrient availability under stress</td>
<td align="left" valign="top"><xref ref-type="bibr" rid="ref94">Offre et al. (2013)</xref>, <xref ref-type="bibr" rid="ref107">Prosser (2012)</xref></td>
</tr>
<tr>
<td align="left" valign="top">Protists</td>
<td align="left" valign="top">Bacterivorous protists</td>
<td align="left" valign="top">Selective bacterivory, regulation of microbial turnover</td>
<td align="left" valign="top">Nutrient mineralization, community succession</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref73">Liao et al. (2024)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Viruses (bacteriophages)</td>
<td align="left" valign="top">Phages</td>
<td align="left" valign="top">Lytic and lysogenic interactions, horizontal gene transfer</td>
<td align="left" valign="top">Microbial population control, functional stability</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref18">Chevallereau et al. (2022)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Microbial BCAs</td>
<td align="left" valign="top"><italic>Trichoderma, Bacillus, Pseudomonas</italic></td>
<td align="left" valign="top">Targeted pathogen suppression, microbiome modulation</td>
<td align="left" valign="top">Disease control, resilience enhancement</td>
<td align="left" valign="top"><xref ref-type="bibr" rid="ref21">Compant et al. (2019)</xref>, <xref ref-type="bibr" rid="ref24">De Faria et al. (2021)</xref></td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="sec5">
<label>2.3</label>
<title>Functional roles in plant health and soil processes</title>
<p>The rhizosphere microbiome contributes to plant health through a set of interrelated functional processes that collectively influence nutrient availability, immune competence, and stress tolerance. Facilitation of nutrient acquisition is a core function, particularly under low-input or stress-prone conditions (<xref ref-type="bibr" rid="ref24">De Faria et al., 2021</xref>). Diverse microbial taxa including <italic>Bacillus</italic>, <italic>Pseudomonas</italic>, <italic>Trichoderma</italic>, and AMF enhance the bioavailability of key nutrients such as phosphorus, potassium, and zinc through mineral solubilization, enzymatic activity, and chelation mechanisms (<xref ref-type="bibr" rid="ref6">Ansabayeva et al., 2025</xref>; <xref ref-type="bibr" rid="ref137">Thepbandit and Athinuwat, 2024</xref>). In parallel, diazotrophic bacteria such as <italic>Azospirillum</italic>, <italic>Burkholderia</italic>, and <italic>Rhizobium</italic> contribute to nitrogen input via biological fixation, supporting plant nutrition in resource-limited systems (<xref ref-type="bibr" rid="ref49">Imran et al., 2021</xref>).</p>
<p>Beyond nutrient-related functions, rhizosphere microorganisms modulate plant physiological responses through hormonal and immune-mediated pathways. Many bacteria and fungi influence plant development by producing or regulating phytohormones, including indole-3-acetic acid (IAA), cytokinins, gibberellins, and ACC deaminase, thereby affecting root architecture and growth dynamics (<xref ref-type="bibr" rid="ref60">Kaya, 2024</xref>; <xref ref-type="bibr" rid="ref95">Orozco-Mosqueda et al., 2023</xref>). These microbial effects are frequently linked to improved water-use efficiency, enhanced nutrient uptake, and increased tolerance to abiotic stress. In addition, specific members of the rhizosphere microbiome can activate induced systemic resistance (ISR) by modulating jasmonic acid (JA) and ethylene (ET) signalling pathways, activating broad-spectrum defense without the growth penalties often associated with systemic acquired resistance (SAR) (<xref ref-type="bibr" rid="ref109">Rabari et al., 2023</xref>; <xref ref-type="bibr" rid="ref156">Zhu et al., 2022</xref>). Well-characterized ISR-inducing taxa include <italic>Pseudomonas fluorescens</italic>, <italic>Bacillus subtilis</italic>, and <italic>Trichoderma harzianum</italic> (<xref ref-type="bibr" rid="ref96">Panpatte et al., 2016</xref>; <xref ref-type="bibr" rid="ref71">Lee D&#x00ED;az et al., 2021</xref>).</p>
<p>The rhizosphere microbiome also contributes to disease suppression through both direct antagonism and community-level interactions. Antagonistic microbes inhibit pathogens via competition for resources, mycoparasitism, and the production of antimicrobial metabolites such as lipopeptides, volatile organic compounds, and hydrolytic enzymes (<xref ref-type="bibr" rid="ref21">Compant et al., 2019</xref>; <xref ref-type="bibr" rid="ref50">Iqbal et al., 2023</xref>). Importantly, pathogen suppression often emerges from the collective activity of stable microbial consortia rather than from individual taxa. Disease-suppressive soils exemplify this principle, as they are consistently associated with enrichment of microbial groups such as <italic>Pseudomonas</italic>, <italic>Trichoderma</italic>, and <italic>Streptomyces</italic>, which constrain pathogens including <italic>Fusarium oxysporum</italic> and <italic>Rhizoctonia solani</italic> through antibiotic production and competition for iron (<xref ref-type="table" rid="tab1">Table 1</xref>; <xref ref-type="bibr" rid="ref24">De Faria et al., 2021</xref>; <xref ref-type="bibr" rid="ref79">Mazzola, 2002</xref>). In addition to biotic interactions, rhizosphere microorganisms play a key role in mitigating abiotic stress. By enhancing antioxidant capacity, osmotic adjustment, and stress-responsive gene expression, beneficial microbes can improve plant tolerance of salinity, drought, and temperature extremes (<xref ref-type="bibr" rid="ref61">Khan et al., 2021</xref>; <xref ref-type="bibr" rid="ref124">Sharma et al., 2020</xref>). However, the magnitude and consistency of these benefits are strongly context-dependent, reflecting interactions between microbial composition, host genotype, and environmental conditions.</p>
</sec>
<sec id="sec6">
<label>2.4</label>
<title>Sensitivity to abiotic and biotic stresses</title>
<p>Despite its ecological adaptability, the rhizosphere microbiome is highly sensitive to environmental disturbance, with abiotic and biotic stressors often interacting to destabilize community structure and function. Abiotic stresses such as drought, salinity, pH fluctuations, and thermal extremes can reduce microbial diversity, suppress keystone taxa, and compromise microbiome-mediated processes that support plant nutrition and defense (<xref ref-type="bibr" rid="ref63">Khan et al., 2023</xref>; <xref ref-type="bibr" rid="ref88">Munir et al., 2022</xref>). For example, drought frequently reduces the abundance of beneficial plant-associated taxa such as <italic>Pseudomonas</italic> and <italic>Bacillus</italic> while favoring desiccation-tolerant groups like Actinobacteria (<xref ref-type="bibr" rid="ref122">Santos-Medell&#x00ED;n et al., 2017</xref>). Similarly, reduced soil moisture availability can limit colonization by AMF, despite its importance for plant water-use efficiency (<xref ref-type="bibr" rid="ref1">Abdalla et al., 2023</xref>). Salinity stress further constrains microbial richness and disrupts symbiotic nutrient acquisition, particularly in non-halophilic taxa, through osmotic and ionic toxicity (<xref ref-type="bibr" rid="ref5">Andronov et al., 2012</xref>; <xref ref-type="bibr" rid="ref88">Munir et al., 2022</xref>).</p>
<p>Biotic stressors, particularly soilborne pathogens, also exert strong selective pressures on rhizosphere communities. Pathogen invasion by species such as <italic>F. oxysporum</italic>, <italic>F. solani</italic>, <italic>Ralstonia solani</italic>, and <italic>R. solani</italic> can restructure microbial networks, suppress beneficial microbes, and induce community dysbiosis (<xref ref-type="table" rid="tab1">Table 1</xref>; <xref ref-type="bibr" rid="ref27">Deng et al., 2021</xref>; <xref ref-type="bibr" rid="ref58">Kashyap et al., 2023</xref>; <xref ref-type="bibr" rid="ref130">Su et al., 2020</xref>). While pathogens may outcompete resident taxa through allelopathy, niche displacement, or immune manipulation, plants can partially counteract these effects through a &#x201C;cry-for-help&#x201D; response, releasing targeted exudates that recruit protective microbes (<xref ref-type="bibr" rid="ref10">Berendsen et al., 2012</xref>; <xref ref-type="bibr" rid="ref117">Rolfe et al., 2019</xref>). This response highlights the dynamic and context-dependent nature of plant&#x2013;microbiome interactions under disease pressure.</p>
<p>Climate change exacerbates the complexity of these interactions by increasing the frequency and intensity of compound stress events such as simultaneous drought and pathogen attack. These multifactorial pressures can undermine both plant immunity and microbiome resilience, thereby reducing the rhizosphere&#x2019;s buffering capacity against further disturbance (<xref ref-type="bibr" rid="ref19">Choudhary and Senthil-Kumar, 2024</xref>). In this context, microbiome resilience reflects the capacity of the rhizosphere community to maintain or rapidly recover key structural and functional attributes following stress. Elevated CO&#x2082; and temperature can further alter root exudation patterns by modifying plant carbon allocation and metabolic activity, reshaping microbial recruitment, and favoring stress-tolerant or opportunistic taxa over beneficial symbionts (<xref ref-type="bibr" rid="ref24">De Faria et al., 2021</xref>; <xref ref-type="bibr" rid="ref72">Li et al., 2021</xref>). Anthropogenic inputs including pesticides and excessive fertilizer treatment may exacerbate these effects by degrading mutualistic networks and simplifying community composition (<xref ref-type="bibr" rid="ref54">Jacobsen and Hjelms&#x00F8;, 2014</xref>). Consequently, sustaining a functionally resilient rhizosphere microbiome depends not only on microbial diversity but also on network redundancy, host compatibility, and the legacy effects of agronomic practices. The development of climate-smart adaptive agroecosystems thus requires knowledge of the thresholds at which beneficial functions collapse and the availability of strategies to restore them through inoculants, amendments, or host breeding.</p>
</sec>
</sec>
<sec id="sec7">
<label>3</label>
<title>Microbial BCAs: key players and mechanisms</title>
<p>Microbial BCAs are a promising class of eco-compatible tools for disease suppression and sustainable crop production. Among the most widely studied genera are <italic>Trichoderma</italic>, <italic>Clonostachys</italic>, <italic>Bacillus</italic>, and <italic>Pseudomonas</italic>, all of which exhibit broad antagonistic activity against phytopathogens and contribute to enhanced plant health under both biotic and abiotic stress (<xref ref-type="bibr" rid="ref40">Guzm&#x00E1;n-Guzm&#x00E1;n et al., 2019</xref>; <xref ref-type="bibr" rid="ref57">Karlsson et al., 2015</xref>; <xref ref-type="bibr" rid="ref69">Lahlali et al., 2022</xref>). While their beneficial effects have mainly been demonstrated under controlled or semi-controlled conditions, these genera exhibit high environmental adaptability and root colonization efficiency, and have multifunctional roles that extend beyond pathogen antagonism.</p>
<p>The core mechanisms by which BCAs act against microbial phytopathogens include antibiosis, ISR, mycoparasitism, and resource competition (<xref ref-type="table" rid="tab2">Table 2</xref>). Antibiosis involves the secretion of antimicrobial metabolites such as lipopeptides, antibiotics, siderophores, and hydrolytic enzymes that inhibit or lyse pathogenic cells. For instance, <italic>B. subtilis</italic> produces surfactins and iturins that disrupt fungal membranes, while <italic>P. fluorescens</italic> is known for its production of phenazine and pyoluteorin compounds with broad-spectrum antifungal activity (<xref ref-type="bibr" rid="ref32">Fischer et al., 2013</xref>). <italic>Trichoderma</italic> species are particularly effective in this regard, releasing cell wall-degrading enzymes such as chitinases and glucanases alongside volatile organic compounds that suppress a wide array of soilborne fungi (<xref ref-type="bibr" rid="ref26">Debode et al., 2018</xref>; <xref ref-type="bibr" rid="ref40">Guzm&#x00E1;n-Guzm&#x00E1;n et al., 2019</xref>). However, the production, stability, and bioavailability of these metabolites are highly context-dependent. These processes are influenced by factors such as soil nutrient status, pH, and competitive pressure from indigenous microbial communities, which often constrains their reproducibility under field conditions.</p>
<table-wrap position="float" id="tab2">
<label>Table 2</label>
<caption>
<p>Representative microbial biocontrol agents (BCAs), their dominant mechanisms of action, and reported effects on rhizosphere microbiome structure and plant performance.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="top">BCA group/genus (examples)</th>
<th align="left" valign="top">Primary mechanisms of action</th>
<th align="left" valign="top">Reported microbiome-level effects</th>
<th align="left" valign="top">Plant/stress-related outcomes</th>
<th align="left" valign="top">References</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="top"><italic>Clonostachys</italic> and <italic>Trichoderma</italic> spp.</td>
<td align="left" valign="top">Mycoparasitism, ISR, enzyme secretion</td>
<td align="left" valign="top">Shifts toward beneficial fungal taxa, increased network connectivity</td>
<td align="left" valign="top">Disease suppression, drought tolerance</td>
<td align="left" valign="top"><xref ref-type="bibr" rid="ref57">Karlsson et al. (2015)</xref>, <xref ref-type="bibr" rid="ref26">Debode et al. (2018)</xref></td>
</tr>
<tr>
<td align="left" valign="top"><italic>Bacillus</italic> spp.</td>
<td align="left" valign="top">Antibiosis, ISR, competition</td>
<td align="left" valign="top">Shifts in bacterial community composition, enrichment of PGPR-associated taxa</td>
<td align="left" valign="top">Pathogen control, improved growth</td>
<td align="left" valign="top"><xref ref-type="bibr" rid="ref21">Compant et al. (2019)</xref>, <xref ref-type="bibr" rid="ref32">Fischer et al. (2013)</xref>, <xref ref-type="bibr" rid="ref109">Rabari et al. (2023)</xref></td>
</tr>
<tr>
<td align="left" valign="top"><italic>Pseudomonas</italic> spp.</td>
<td align="left" valign="top">Antibiotic production, siderophores, ISR</td>
<td align="left" valign="top">Suppression of pathogens, altered rhizosphere structure</td>
<td align="left" valign="top">Disease resistance, nutrient uptake</td>
<td align="left" valign="top"><xref ref-type="bibr" rid="ref69">Lahlali et al. (2022)</xref>, <xref ref-type="bibr" rid="ref79">Mazzola (2002)</xref></td>
</tr>
<tr>
<td align="left" valign="top">Microbial consortia</td>
<td align="left" valign="top">Functional complementarity among microbial members</td>
<td align="left" valign="top">Increased microbial diversity and stability</td>
<td align="left" valign="top">Enhanced stress resilience</td>
<td align="left" valign="top"><xref ref-type="bibr" rid="ref143">Vishwakarma et al. (2020)</xref>, <xref ref-type="bibr" rid="ref75">Mamun et al. (2024)</xref></td>
</tr>
</tbody>
</table>
</table-wrap>
<p>Mycoparasitism, a direct parasitic interaction in which a biocontrol fungus invades and degrades a pathogenic one, is well-documented in <italic>T. harzianum</italic> and <italic>Clonostachys rosea</italic> (<xref ref-type="bibr" rid="ref17">Chet, 1987</xref>; <xref ref-type="bibr" rid="ref57">Karlsson et al., 2015</xref>). These fungi recognize pathogen-specific molecular patterns, initiate targeted enzymatic degradation, and physically coil around or penetrate pathogen hyphae, leading to pathogen death. This process is often coupled with competition for space and nutrients in the rhizosphere, effectively excluding pathogens from becoming established on the root surface. Despite its favorable effects in controlled systems, effective mycoparasitism in dynamic soil environments requires the establishment and persistence of the mycoparasitic agent in the soil, which is often difficult to achieve with introduced BCAs. In addition to their antagonistic functions, some BCAs play an important role in priming plant defense responses. Many strains induce ISR, a plant-mediated immune enhancement triggered by microbial signals such as lipopolysaccharides or flagellins (<xref ref-type="bibr" rid="ref103">Pieterse et al., 2014</xref>). Unlike SAR, ISR typically does not involve accumulation of salicylic acid (SA) or pathogenesis-related (PR) proteins, allowing plants to maintain growth while remaining primed for faster and stronger defense upon pathogen attack (<xref ref-type="bibr" rid="ref44">Heil and Bostock, 2002</xref>). Emerging evidence also highlights the role of microbial BCAs in improving plant tolerance to abiotic stress, a less explored but increasingly relevant function under climate change scenarios. For example, <italic>Trichoderma</italic> spp. can improve water-use efficiency, enhance antioxidant activity, and modulate stress-responsive genes, thereby mitigating the adverse effects of drought and salinity (<xref ref-type="bibr" rid="ref127">Sorahinobar et al., 2025</xref>). Similarly, <italic>Bacillus</italic> spp. have been shown to promote root growth and osmolyte accumulation under heat and osmotic stress conditions (<xref ref-type="bibr" rid="ref92">Nivetha et al., 2024</xref>). However, much of this evidence derives from laboratory or greenhouse experiments, and the desired beneficial responses are not consistently observed in heterogeneous field environments. Further study is thus needed to clarify the molecular and physiological pathways underpinning these benefits in complex, multi-stress field environments, and to evaluate their quantitative impacts on yields under severe stress.</p>
</sec>
<sec id="sec8">
<label>4</label>
<title>Interactions between microbial BCAs and the rhizosphere microbiome</title>
<sec id="sec9">
<label>4.1</label>
<title>Microbiome modulation by microbial BCAs</title>
<p>The introduction of microbial BCAs into the rhizosphere to suppress phytopathogens also modulates native microbial communities via both direct and indirect mechanisms that influence community interactions and stability. These mechanisms include niche competition, secretion of antibiotics, interference with microbial signalling pathways, and metabolic cross-feeding, wherein one microbe utilizes metabolites produced by another (<xref ref-type="table" rid="tab2">Table 2</xref>; <xref ref-type="bibr" rid="ref135">Tang et al., 2025</xref>; <xref ref-type="bibr" rid="ref153">Zhang et al., 2023</xref>). In addition to these direct microbial interactions, BCAs can influence rhizosphere community assembly through plant-mediated effects. BCA inoculation has been shown to modify the volume and composition of root exudates, for example by changing the content of sugars, organic acids, amino acids, and secondary metabolites, which in turn shape microbial recruitment and activity in the rhizosphere. Such exudate-mediated shifts can selectively promote beneficial taxa or alter competitive interactions among resident microbes, thereby indirectly contributing to microbiome restructuring and functional outcomes (<xref ref-type="bibr" rid="ref10">Berendsen et al., 2012</xref>; <xref ref-type="bibr" rid="ref41">Haichar et al., 2008</xref>; <xref ref-type="bibr" rid="ref117">Rolfe et al., 2019</xref>). Through these processes, BCAs influence microbial diversity, including both alpha diversity (within-sample richness) and beta diversity (between-sample compositional differences). The magnitude and direction of these shifts depend on the host plant and soil type as well as the resident microbiota (<xref ref-type="bibr" rid="ref26">Debode et al., 2018</xref>; <xref ref-type="bibr" rid="ref147">Wilson and Arunachalam, 2024</xref>). Inoculation with BCAs can thus act as an ecological perturbation, triggering shifts in community structure and ecosystem function. However, such modulation is not universally beneficial. In cases of niche overlap or ecological antagonism, BCAs may competitively displace native microbial taxa crucial for nutrient cycling or stress buffering. This can potentially trigger cascading nontarget effects across the rhizosphere microbiome (<xref ref-type="bibr" rid="ref99">Pearson and Callaway, 2003</xref>, <xref ref-type="bibr" rid="ref100">2005</xref>; <xref ref-type="bibr" rid="ref138">Trabelsi and Mhamdi, 2013</xref>). Additionally, BCAs fail to persist in some settings due to poor compatibility with the indigenous microbiome or unfavorable soil conditions, reducing their long-term efficacy (<xref ref-type="bibr" rid="ref123">Schulz et al., 2019</xref>).</p>
</sec>
<sec id="sec10">
<label>4.2</label>
<title>Factors influencing inoculant performance</title>
<p>The effectiveness of BCAs in shaping rhizosphere microbiomes is highly variable, particularly when comparing laboratory, greenhouse, and field settings, reflecting the interplay of abiotic, biotic, and ecological factors. Soil physicochemical traits such as pH, texture, nutrient availability, and organic matter content are key abiotic drivers that strongly influence microbial colonization and functional expression (<xref ref-type="bibr" rid="ref12">Berg and Smalla, 2009</xref>; <xref ref-type="bibr" rid="ref138">Trabelsi and Mhamdi, 2013</xref>). For instance, acidic soils can limit <italic>Bacillus</italic> spp. sporulation, while organic-rich soils may reduce the competitive advantage of nutrient-solubilizing strains. Beyond effects on colonization, abiotic conditions can directly regulate the expression of BCA functions including the biosynthesis and stability of antibiotics and other secondary metabolites, thereby modulating antagonistic activity independently of population size (<xref ref-type="bibr" rid="ref12">Berg and Smalla, 2009</xref>; <xref ref-type="bibr" rid="ref108">Raaijmakers and Mazzola, 2012</xref>). Such context-specific responses underscore the challenge of developing universally effective formulations.</p>
<p>Plant genotype exerts an equally strong filtering effect. Root exudate chemistry, architectural traits, and immune signalling not only shape microbial recruitment but also determine compatibility with inoculated strains. Genotype-specific filtering can facilitate or block BCA establishment depending on how introduced microbes overlap with resident microbiota niches (<xref ref-type="bibr" rid="ref102">Philippot et al., 2013</xref>; <xref ref-type="bibr" rid="ref111">Rawal et al., 2024</xref>). This highlights the need to consider crop genetic background in inoculant design, particularly in breeding programs that historically overlook plant&#x2013;microbe interactions (<xref ref-type="bibr" rid="ref51">Iqbal et al., 2025</xref>). Climatic variability adds another layer of complexity. Temperature extremes, soil moisture fluctuations, and seasonal cycles modulate microbial persistence and activity. For example, <italic>B. subtilis</italic> and <italic>T. harzianum</italic> have shown strong antagonism in controlled trials but exhibit inconsistent or diminished efficacy under field conditions, suggesting that environmental complexity often limits the translation of laboratory efficacy (<xref ref-type="bibr" rid="ref113">Rigobelo et al., 2024</xref>).</p>
<p>Colonization and persistence are critical bottlenecks. Many inoculants fail to establish durable populations due to competitive exclusion by native microbiota, predation by protists, or suboptimal timing and delivery methods. In addition to effects on establishment, interactions with indigenous microbial communities can directly constrain or enhance BCA efficacy through interspecies metabolic interference or facilitation. For example, <italic>Stenotrophomonas maltophilia</italic> can degrade lipopeptide antibiotics produced by <italic>Bacillus</italic> spp. (e.g., iturin, fengycin, surfactin), thereby attenuating biocontrol of tomato bacterial wilt despite successful BCA colonization (<xref ref-type="bibr" rid="ref101">Peng et al., 2025</xref>). In contrast, positive interactions with resident microbial keystone taxa can enhance BCA performance. A recent study showed that specific indigenous microbes stimulated antibiotic biosynthesis in <italic>Streptomyces</italic>, leading to improved disease suppression through metabolite-mediated induction rather than direct antagonism alone (<xref ref-type="bibr" rid="ref133">Sun et al., 2025</xref>). These contrasting outcomes highlight that BCA efficacy is highly context-dependent rather than universally reproducible. Even when initial colonization is achieved, BCA-mediated functions such as ISR or mycoparasitism are often regulated by density-dependent microbial signalling and plant-derived cues, which may not be consistently activated across diverse soil environments (<xref ref-type="bibr" rid="ref86">Moussa and Iasur Kruh, 2025</xref>). Moreover, traits observed during <italic>in vitro</italic> screening often fail to translate to complex rhizosphere environments, where multi-trophic interactions, fluctuating nutrient availability, and stress events shape microbial behavior. This discrepancy illustrates the limitations of current screening pipelines, which often neglect ecological compatibility and stress resilience.</p>
</sec>
<sec id="sec11">
<label>4.3</label>
<title>Risks and trade-offs</title>
<p>Despite their potential, BCAs are not without risks. One critical concern is the displacement of native beneficial microbes, particularly in systems with already well-functioning microbial communities. The introduction of dominant inoculants can reduce microbial diversity or suppress indigenous taxa that play critical roles in nutrient cycling, pathogen suppression, and stress buffering (<xref ref-type="bibr" rid="ref99">Pearson and Callaway, 2003</xref>; <xref ref-type="bibr" rid="ref138">Trabelsi and Mhamdi, 2013</xref>). Such non-target effects may compromise essential ecosystem functions if introduced BCAs competitively displace resident microbes that are functionally important but not directly associated with disease suppression. These ecological trade-offs are especially evident in monospecific inoculations, which often lack the resilience and functional redundancy characteristic of native microbial consortia. Many BCAs also exhibit narrow-spectrum efficacy, performing effectively against specific pathogens under controlled conditions but showing limited broad-spectrum protection and consistency in complex field environments (<xref ref-type="bibr" rid="ref70">Leal et al., 2024</xref>).</p>
<p>Beyond these general constraints, more subtle ecological risks may arise. Targeting a single pathogen can inadvertently lead to ecological release of secondary or opportunistic pathogens, thereby unpredictably altering disease complexes. Unintended effects may also result from horizontal gene transfer, interference with microbial signalling networks, or suppression of keystone taxa that support the rhizosphere&#x2019;s structural stability. Repeated or high-frequency BCA applications may amplify these effects by cumulatively reshaping microbial networks and selecting for altered interaction dynamics over time. Such perturbations can compromise microbiome resilience, reduce invasion resistance, and impair ecosystem services that underpin long-term soil health. Importantly, these risks are context-dependent: BCAs may enhance resilience by restoring lost functions in degraded or simplified systems but could disrupt established microbial equilibria in biodiverse or organically managed soils. Despite these concerns, there have been few long-term evaluations of the ecological consequences of repeated BCA applications across cropping cycles and environmental conditions. Current risk assessment approaches are largely oriented toward short-term efficacy, with less emphasis on microbiome-level impacts, biosafety considerations, or cumulative ecological effects, highlighting the need for more holistic and longitudinal perspectives on the sustainability of microbial biocontrol interventions.</p>
</sec>
</sec>
<sec id="sec12">
<label>5</label>
<title>Climate-responsive microbial biocontrol</title>
<sec id="sec13">
<label>5.1</label>
<title>Fungal pathogens and climate change</title>
<p>Climate change is reshaping the global epidemiology of fungal diseases, with increasing incidences attributed to drought, salinity, temperature extremes, and shifting precipitation patterns (<xref ref-type="bibr" rid="ref9">Bebber et al., 2013</xref>; <xref ref-type="bibr" rid="ref126">Singh et al., 2023</xref>). These stressors extend the geographic range of pathogens such as <italic>Fusarium</italic> spp., <italic>Verticillium</italic> spp., and <italic>Botrytis cinerea</italic>, while simultaneously enhancing their aggressiveness and survival under adverse conditions.</p>
<p>Plant susceptibility is aggravated under abiotic stress. Drought impairs root development, alters rhizodeposition patterns, and limits the plant&#x2019;s ability to recruit beneficial microbes. These factors collectively increase plants&#x2019; vulnerability to soilborne pathogens (<xref ref-type="bibr" rid="ref106">Preece and Pe&#x00F1;uelas, 2016</xref>; <xref ref-type="bibr" rid="ref150">Zahra et al., 2023</xref>). Likewise, salinity and high temperatures have been shown to downregulate key immune signalling pathways, including those regulated by SA, JA, and ET, which are essential for SAR and induced defense responses (<xref ref-type="bibr" rid="ref118">Rossi et al., 2023</xref>). Importantly, interactions between abiotic and biotic stressors are often synergistic rather than additive. Combined stress conditions, such as drought coinciding with a pathogen challenge, can lead to greater physiological damage than either stress alone. This is partly due to compounded suppression of plant immune responses and microbiome-mediated defense mechanisms. These interactions also reduce the functional stability of rhizosphere microbial communities, which limits their capacity to buffer against pathogen invasion (<xref ref-type="bibr" rid="ref110">Ramegowda and Senthil-Kumar, 2015</xref>; <xref ref-type="bibr" rid="ref134">Suzuki et al., 2014</xref>).</p>
</sec>
<sec id="sec14">
<label>5.2</label>
<title>Microbial BCA strategies for climate resilience</title>
<p>Microbial BCAs provide two key functions under climate stress conditions: they suppress phytopathogens while simultaneously enhancing plant tolerance of abiotic stressors (<xref ref-type="bibr" rid="ref125">Sheoran et al., 2025</xref>). These functions are mediated by stressor-specific mechanisms that operate at both the cellular and rhizosphere ecosystem levels and are linked to key climate variables including drought-induced water limitation, temperature-driven metabolic constraints, and salinity-associated osmotic stress. However, these benefits are not consistently observed across experimental scales: desirable outcomes are seen more frequently under controlled conditions than in heterogeneous field environments. Understanding and optimizing these interactions will be essential for developing climate-resilient cropping systems. Representative studies demonstrating BCA-induced microbiome and plant responses under stress conditions are summarized in <xref ref-type="table" rid="tab3">Table 3</xref>.</p>
<table-wrap position="float" id="tab3">
<label>Table 3</label>
<caption>
<p>Representative studies demonstrating microbial biocontrol agents (BCAs)&#x2013;induced changes in microbiome structure and plant responses under stress conditions relevant to climate resilience.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="top">Stress condition</th>
<th align="left" valign="top">BCA system (example)</th>
<th align="left" valign="top">Observed microbiome/plant response</th>
<th align="left" valign="top">Level of response</th>
<th align="left" valign="top">References</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="top">Drought</td>
<td align="left" valign="top"><italic>Trichoderma</italic> spp.</td>
<td align="left" valign="top">Enrichment of stress-tolerant taxa; enhanced antioxidant activity</td>
<td align="left" valign="top">Microbiome + plant</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref85">Mona et al. (2017)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Salinity</td>
<td align="left" valign="top">PGPR (<italic>Bacillus</italic>, <italic>Pseudomonas</italic>)</td>
<td align="left" valign="top">Osmolyte accumulation; improved ion homeostasis</td>
<td align="left" valign="top">Plant physiology</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref43">Ha-Tran et al. (2021)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Drought/salinity</td>
<td align="left" valign="top">PGPR</td>
<td align="left" valign="top">Microbial community restructuring; improved yield stability</td>
<td align="left" valign="top">Microbiome + plant</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref4">Al-Turki et al. (2023)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Multi-stress</td>
<td align="left" valign="top">Microbial consortia</td>
<td align="left" valign="top">Increased community stability and functional redundancy</td>
<td align="left" valign="top">Microbiome</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref75">Mamun et al. (2024)</xref>
</td>
</tr>
</tbody>
</table>
</table-wrap>
<sec id="sec15">
<label>5.2.1</label>
<title>Stress-responsive modulation by BCAs</title>
<p>Fungal BCAs, particularly <italic>Trichoderma</italic> spp., have a well-documented capacity to mitigate drought- and heat-associated stress through coordinated modulation of plant antioxidant systems, osmotic balance, and hormonal pathways (<xref ref-type="bibr" rid="ref36">Geng et al., 2025</xref>). In maize and tomato, inoculation with <italic>T. asperellum</italic> and <italic>T. harzianum</italic> significantly enhanced the expression and activity of key antioxidant enzymes such as superoxide dismutase (SOD), catalase (CAT), and ascorbate peroxidase (APX), which all play important roles in detoxifying reactive oxygen species (ROS) generated under water-deficit conditions (<xref ref-type="bibr" rid="ref85">Mona et al., 2017</xref>; <xref ref-type="bibr" rid="ref149">Yousaf et al., 2023</xref>). Such biochemical priming reduces oxidative cellular damage, stabilizes membrane integrity, and improves photosynthetic efficiency primarily in laboratory and greenhouse studies. Moreover, plant growth-promoting rhizobacteria (PGPR) such as <italic>P. fluorescens</italic>, <italic>Azospirillum brasilense</italic>, and <italic>B. subtilis</italic> have demonstrated the capacity to improve plant performance under drought- and salinity-driven osmotic stress (<xref ref-type="table" rid="tab3">Table 3</xref>). These bacteria modulate the synthesis and accumulation of osmolytes such as proline, trehalose, and glycine betaine, which facilitate cellular osmotic adjustment and help maintain turgor pressure under dehydrating conditions (<xref ref-type="bibr" rid="ref43">Ha-Tran et al., 2021</xref>). Several PGPR strains produce exopolysaccharides (EPS) that enhance soil aggregation and improve water-holding capacity, partially offsetting moisture limitation in drought-prone soils (<xref ref-type="bibr" rid="ref155">Zheng et al., 2018</xref>). Moreover, some strains trigger phytohormonal crosstalk by producing analogues or regulators of IAA, abscisic acid (ABA), and ET, thereby influencing root system architecture and stomatal conductance (<xref ref-type="bibr" rid="ref62">Khan et al., 2020</xref>). This hormonal tuning facilitates nutrient uptake under stress and also contributes to adaptive regulation of plant water use under variable climatic conditions. These microbial mechanisms represent promising bio-based tools for buffering crops against increasingly frequent climate-induced stressors.</p>
</sec>
<sec id="sec16">
<label>5.2.2</label>
<title>Mechanistic insights from omics technologies</title>
<p>Advances in omics technologies have provided valuable insights into the mechanistic basis of microbe-mediated stress tolerance in plants under specific climate-related stressors. Transcriptomic analyses of plants inoculated with <italic>Trichoderma</italic> spp. revealed the upregulation of stress-responsive genes involved in aquaporin regulation, expression of late embryogenesis abundant (LEA) proteins, and ABA signalling, among other things. These changes collectively contribute to improved water retention, membrane protection, and stomatal regulation under drought conditions (<xref ref-type="bibr" rid="ref30">Doni et al., 2019</xref>; <xref ref-type="bibr" rid="ref154">Zhang et al., 2022</xref>). At the molecular level, these adjustments show how fungal BCAs can fine-tune host stress physiology in addition to performing their traditional biocontrol roles. Metabolomic studies have similarly shown that PGPR elicit the synthesis of osmo-protectants such as proline and glycine betaine as well as secondary metabolites such as flavonoids and phenolics that function as ROS scavengers and cell protectants during abiotic stress (<xref ref-type="bibr" rid="ref4">Al-Turki et al., 2023</xref>). In salt-stressed tomato and soybean plants, PGPR-induced metabolic reprogramming has been linked to improved ion homeostasis, photosynthetic stability, and yield retention (<xref ref-type="bibr" rid="ref4">Al-Turki et al., 2023</xref>; <xref ref-type="bibr" rid="ref68">Kumawat et al., 2023</xref>). Amplicon-based microbiome profiling (16S rRNA and ITS sequencing) has demonstrated that BCAs can also restructure rhizosphere microbial networks, promoting the enrichment of drought-resilient taxa such as Actinobacteria and certain classes of Proteobacteria while reducing the relative abundance of opportunistic pathogens (<xref ref-type="bibr" rid="ref35">Gao et al., 2023</xref>). These shifts reflect functional selection for stress-tolerant guilds, potentially enhancing microbial synergy and functional redundancy under environmental perturbations. However, it should be noted that most omics-based insights derive from controlled systems, and their predictive value for field-scale resilience remains limited.</p>
</sec>
<sec id="sec17">
<label>5.2.3</label>
<title>Role of microbial consortia and formulations</title>
<p>Microbial consortia are mixtures of multiple co-inoculated bacterial and/or fungal strains. It is increasingly recognized that such consortia can offer greater stability and multifunctionality than single strain inoculations under climate-induced stress conditions. Unlike monospecific BCAs, consortia can exploit functional complementarity wherein different microbes perform distinct but synergistic roles such as nutrient solubilization, pathogen suppression, and hormone modulation (<xref ref-type="bibr" rid="ref143">Vishwakarma et al., 2020</xref>). The combined application of <italic>T. harzianum</italic>, <italic>B. subtilis</italic>, and AMF in tomato, strawberry, and potato systems enhanced drought tolerance and disease suppression, which was attributed to co-regulation of ABA signalling, antioxidant enzyme activation, and root colonization efficiency (<xref ref-type="bibr" rid="ref75">Mamun et al., 2024</xref>; <xref ref-type="bibr" rid="ref83">Minchev et al., 2021</xref>). However, consistent performance of consortia is not guaranteed: antagonism among consortium members or competition with resident microbiota can negate expected benefits if ecological compatibility is not adequately assessed. Formulation technologies are another important determinant of field efficacy. Encapsulation in biopolymeric matrices such as alginate beads, biochar, or nanocarriers has been shown to enhance microbial viability under environmental stress by buffering against UV radiation, desiccation, and oxidative stress (<xref ref-type="bibr" rid="ref8">Balla et al., 2022</xref>; <xref ref-type="bibr" rid="ref22">Cruz-Barrera et al., 2024</xref>). Even so, formulation success remains highly context-dependent and delivery systems must be tailored to specific soil textures, pH ranges, and moisture regimes in order to optimize BCA release kinetics and colonization dynamics in climate-challenged regions.</p>
</sec>
</sec>
</sec>
<sec id="sec18">
<label>6</label>
<title>Case studies in crop systems: microbiome and biocontrol interactions</title>
<p>The following case studies illustrate how interactions between microbial BCAs and the rhizosphere microbiome manifest across diverse crop systems and stress contexts. While each example highlights crop-specific outcomes, they collectively reveal recurrent patterns in microbiome modulation, alongside pronounced context dependence driven by host traits, environmental conditions, and management practices. <xref ref-type="table" rid="tab4">Table 4</xref> presents a synthesis comparing the features of these systems and highlighting key divergences.</p>
<table-wrap position="float" id="tab4">
<label>Table 4</label>
<caption>
<p>Comparative synthesis of microbial biocontrol agents (BCAs)&#x2013;microbiome interactions across crop systems.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="top">Crop system</th>
<th align="left" valign="top">Representative BCAs</th>
<th align="left" valign="top">Dominant stress context</th>
<th align="left" valign="top">Microbiome-level effects</th>
<th align="left" valign="top">Plant-level outcomes</th>
<th align="left" valign="top">Key sources of variability</th>
<th align="left" valign="top">References</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="top">Strawberry</td>
<td align="left" valign="top"><italic>Trichoderma</italic>, <italic>Bacillus</italic>, <italic>Aureobasidium</italic></td>
<td align="left" valign="top">Soilborne pathogens; postharvest stress</td>
<td align="left" valign="top">Increased diversity, enhanced network connectivity, enrichment of keystone taxa</td>
<td align="left" valign="top">Disease suppression, improved fruit quality</td>
<td align="left" valign="top">Baseline microbiome composition, delivery method</td>
<td align="left" valign="top"><xref ref-type="bibr" rid="ref26">Debode et al. (2018)</xref>, <xref ref-type="bibr" rid="ref52">Iqbal et al. (2021)</xref>, <xref ref-type="bibr" rid="ref56">Kang et al. (2025)</xref>, <xref ref-type="bibr" rid="ref81">Men&#x00E9;ndez-Ca&#x00F1;amares et al. (2024)</xref></td>
</tr>
<tr>
<td align="left" valign="top">Tomato/Cucurbits</td>
<td align="left" valign="top"><italic>Bacillus</italic>, <italic>Pseudomonas</italic>, <italic>Trichoderma</italic></td>
<td align="left" valign="top">Pathogen pressure; drought; salinity</td>
<td align="left" valign="top">Shifts in bacterial&#x2013;fungal balance, AMF enrichment</td>
<td align="left" valign="top">Disease control, stress tolerance</td>
<td align="left" valign="top">Host genotype, soil type, stress intensity</td>
<td align="left" valign="top"><xref ref-type="bibr" rid="ref2">Ali et al. (2025)</xref>, <xref ref-type="bibr" rid="ref80">Mehmood et al. (2023)</xref>, <xref ref-type="bibr" rid="ref86">Moussa and Iasur Kruh (2025)</xref></td>
</tr>
<tr>
<td align="left" valign="top">Grapevine/Banana</td>
<td align="left" valign="top"><italic>Trichoderma</italic>, <italic>Bacillus</italic></td>
<td align="left" valign="top">Chronic pathogen pressure; drought</td>
<td align="left" valign="top">Increased functional redundancy, endosphere restructuring</td>
<td align="left" valign="top">Reduced disease severity, yield stability</td>
<td align="left" valign="top">Crop perenniality, climate variability</td>
<td align="left" valign="top"><xref ref-type="bibr" rid="ref16">Chen et al. (2024)</xref>, <xref ref-type="bibr" rid="ref23">Damodaran et al. (2020)</xref>, <xref ref-type="bibr" rid="ref29">Di Marco et al. (2004)</xref></td>
</tr>
<tr>
<td align="left" valign="top">Rice/Maize</td>
<td align="left" valign="top">Microbial consortia (N-fixers, P-solubilizers, BCAs)</td>
<td align="left" valign="top">Salinity; temperature stress</td>
<td align="left" valign="top">Functional complementarity, enhanced nutrient cycling</td>
<td align="left" valign="top">Biomass and yield maintenance</td>
<td align="left" valign="top">Cultivar compatibility, land-use history</td>
<td align="left" valign="top"><xref ref-type="bibr" rid="ref77">Marzouk et al. (2025)</xref>, <xref ref-type="bibr" rid="ref119">Sackey et al. (2025)</xref>, <xref ref-type="bibr" rid="ref140">Tyagi et al. (2023)</xref></td>
</tr>
</tbody>
</table>
</table-wrap>
<sec id="sec19">
<label>6.1</label>
<title>Strawberry as a model system</title>
<p>Strawberry (<italic>Fragaria</italic> &#x00D7; <italic>ananassa</italic>) has emerged as a valuable model for studying interactions between microbial BCAs and rhizosphere microbiome dynamics due to its relatively short lifecycle, well-characterized microbial baseline, and high sensitivity to biotic and abiotic stresses (<xref ref-type="bibr" rid="ref26">Debode et al., 2018</xref>; <xref ref-type="bibr" rid="ref56">Kang et al., 2025</xref>). Its vulnerability to soilborne pathogens including <italic>B. cinerea</italic>, <italic>F. oxysporum</italic>, and <italic>Colletotrichum acutatum</italic>, combined with a microbiome that is both diverse and responsive to disturbance, provides a high-resolution framework for testing microbiome-modulating interventions (<xref ref-type="bibr" rid="ref56">Kang et al., 2025</xref>). Inoculation with BCAs such as <italic>T. harzianum</italic>, <italic>Aureobasidium pullulans</italic>, and <italic>B. subtilis</italic> have consistently suppressed <italic>B. cinerea</italic> and <italic>C. acutatum</italic> while simultaneously enriching rhizosphere diversity and microbial network connectivity in strawberry studies (<xref ref-type="bibr" rid="ref34">Freeman et al., 2004</xref>; <xref ref-type="bibr" rid="ref47">Huang et al., 2023</xref>; <xref ref-type="bibr" rid="ref52">Iqbal et al., 2021</xref>, <xref ref-type="bibr" rid="ref50">2023</xref>; <xref ref-type="bibr" rid="ref76">Mannaa et al., 2023</xref>; <xref ref-type="bibr" rid="ref81">Men&#x00E9;ndez-Ca&#x00F1;amares et al., 2024</xref>). These BCAs suppress pathogens via both indirect antagonistic mechanisms, including antibiosis, and direct interactions such as mycoparasitism. They also appear to facilitate beneficial shifts in core microbial taxa associated with nutrient mobilization, systemic resistance, and root colonization. Bee-vectored delivery of <italic>C. rosea</italic> and <italic>A. pullulans</italic> has demonstrated dual benefits, enhancing pre-harvest disease resistance and improving post-harvest fruit quality by extending shelf life and reducing decay incidence (<xref ref-type="bibr" rid="ref53">Iqbal et al., 2022</xref>; <xref ref-type="bibr" rid="ref67">Kumar et al., 2024</xref>). These strategies showcase systems-level integration of biocontrol, highlighting its potential application across plant developmental stages and the agro-supply chain. Importantly, in strawberry systems, BCA inoculation has been shown to alter the rhizosphere fungal community beyond direct pathogen suppression, including changes in community composition and associated plant defense responses (<xref ref-type="bibr" rid="ref26">Debode et al., 2018</xref>).</p>
</sec>
<sec id="sec20">
<label>6.2</label>
<title>Tomato, muskmelon, and cucumber: managing pathogen pressure and abiotic stress</title>
<p>Tomato (<italic>Solanum lycopersicum</italic>), muskmelon (<italic>Cucumis melo</italic>), and cucumber (<italic>Cucumis sativus</italic>) have been important systems for evaluating the dual role of microbial BCAs in disease suppression and abiotic stress mitigation. In tomato, complex microbial consortia comprising <italic>P. fluorescens</italic>, <italic>T. asperellum</italic>, <italic>B. velezensis</italic>, and <italic>B. subtilis</italic> have shown efficacy against a spectrum of soilborne pathogens, including <italic>R. solanacearum</italic>, <italic>F. oxysporum</italic> f.sp. <italic>lycopersici</italic>, and <italic>Meloidogyne</italic> spp. (root-knot nematodes) (<xref ref-type="bibr" rid="ref2">Ali et al., 2025</xref>; <xref ref-type="bibr" rid="ref39">Gu et al., 2023</xref>; <xref ref-type="bibr" rid="ref86">Moussa and Iasur Kruh, 2025</xref>; <xref ref-type="bibr" rid="ref93">Obiazikwor et al., 2025</xref>; <xref ref-type="bibr" rid="ref145">Wan et al., 2017</xref>). These BCAs not only provided biocontrol through direct antagonism and ISR induction but also reshaped the microbial composition of the rhizosphere, shifting patterns of alpha and beta diversity. Importantly, BCA application was associated with a higher abundance of Actinobacteria and mycorrhizal fungi such as <italic>Glomus</italic> spp., which are known to contribute to enhanced phosphorus solubilization, root colonization, and pathogen exclusion (<xref ref-type="bibr" rid="ref39">Gu et al., 2023</xref>; <xref ref-type="bibr" rid="ref145">Wan et al., 2017</xref>).</p>
<p>Promising results have also been obtained in cucurbit systems, particularly muskmelon and cucumber. Applications of <italic>B. amyloliquefaciens</italic>, <italic>B. subtilis</italic>, and <italic>Pseudomonas chlororaphis</italic> reduced the incidence of <italic>F. oxysporum</italic> f. sp. <italic>radicis-cucumerinum</italic>, <italic>Podosphaera xanthii</italic>, <italic>B. cinerea</italic>, <italic>R. solani</italic> and <italic>Pythium</italic> spp., especially under drought and saline conditions in which chemical controls often fail (<xref ref-type="bibr" rid="ref80">Mehmood et al., 2023</xref>; <xref ref-type="bibr" rid="ref90">Ni and Punja, 2020</xref>). The BCAs also activated host stress responses including the accumulation of antioxidant enzymes (e.g., SOD, CAT, APX) and compatible solutes (e.g., proline, glycine betaine), thus contributing to osmotic adjustment and oxidative damage mitigation. These results emphasize that effective biocontrol is not limited to pathogen suppression but often coincides with improved rhizosphere functionality and resilience.</p>
</sec>
<sec id="sec21">
<label>6.3</label>
<title>Grapevine and banana: trunk disease and wilt suppression</title>
<p>Perennial crops such as grapevine (<italic>Vitis vinifera</italic>) and banana (<italic>Musa</italic> spp.) are high-value systems that are increasingly threatened by persistent fungal pathogens and environmental stressors. In grapevines, trunk diseases caused by <italic>Phaeomoniella chlamydospora</italic> and <italic>Eutypa lata</italic> have shown significant responsiveness to treatment with <italic>T. atroviride</italic>, <italic>T. harzianum</italic> and <italic>B. subtilis</italic>, especially in Mediterranean climates characterized by thermal variability and drought stress (<xref ref-type="bibr" rid="ref29">Di Marco et al., 2004</xref>; <xref ref-type="bibr" rid="ref38">Gramaje et al., 2018</xref>; <xref ref-type="bibr" rid="ref65">Kotze et al., 2011</xref>; <xref ref-type="bibr" rid="ref82">Mesguida et al., 2023</xref>). These BCAs reduced disease severity while also inducing restructuring of both rhizosphere and endosphere microbial communities. This led to increased functional redundancy in bacterial guilds associated with xenobiotic degradation, biosynthesis of phytohormones including auxins and gibberellins, and immune modulation, suggesting enhanced ecological stability under pathogen pressure (<xref ref-type="bibr" rid="ref16">Chen et al., 2024</xref>).</p>
<p>In bananas, inoculation with <italic>T. asperellum</italic>, <italic>T. reesei</italic>, and <italic>T. koningiopsis</italic> achieved partial control of <italic>F. oxysporum</italic> f. sp. <italic>cubense</italic> tropical race 4 (Foc TR4), a pathogen of global concern (<xref ref-type="bibr" rid="ref23">Damodaran et al., 2020</xref>; <xref ref-type="bibr" rid="ref104">Ploetz, 2015</xref>). Field studies across Asia and Latin America showed consistent suppression of disease symptoms alongside improved yield metrics. Treated plants exhibited elevated expression of PR-related genes, improved root architecture, and an increase in rhizospheric populations of plant growth-promoting bacteria (PGPB), contributing to both biotic resistance and physiological resilience (<xref ref-type="bibr" rid="ref84">Mon et al., 2021</xref>).</p>
</sec>
<sec id="sec22">
<label>6.4</label>
<title>Rice, maize, and other staples: consortia and microbiome modulation</title>
<p>Cereal staples such as rice (<italic>Oryza sativa</italic>) and maize (<italic>Zea mays</italic>) are increasingly being targeted for treatment with microbial consortia to enhance stress resilience and nutrient efficiency under climate stress (<xref ref-type="bibr" rid="ref55">Jha et al., 2013</xref>; <xref ref-type="bibr" rid="ref140">Tyagi et al., 2023</xref>; <xref ref-type="bibr" rid="ref152">Zayadan et al., 2014</xref>). These consortia typically combine functionally complementary taxa like nitrogen-fixers (<italic>Azospirillum</italic>, <italic>Serratia</italic>), phosphate-solubilizers (<italic>Azotobacter, Bacillus</italic>, <italic>Enterobacter</italic>), and biocontrol fungi (<italic>Trichoderma</italic> spp.) to simultaneously address multiple agroecological constraints (<xref ref-type="bibr" rid="ref3">Altomare et al., 1999</xref>; <xref ref-type="bibr" rid="ref25">De Vries and Wallenstein, 2017</xref>; <xref ref-type="bibr" rid="ref45">Hermosa et al., 2013</xref>; <xref ref-type="bibr" rid="ref64">Kong et al., 2018</xref>; <xref ref-type="bibr" rid="ref115">Rizvi and Khan, 2018</xref>; <xref ref-type="bibr" rid="ref148">Woo and Pepe, 2018</xref>). Under both controlled and field environment, such inoculants have been shown to improve crop performance, particularly in low-input systems exposed to salinity or thermal extremes. Mechanistically, these benefits are linked to enhanced ion homeostasis (e.g., improved K<sup>+</sup>/Na<sup>+</sup> ratios), modulation of antioxidant enzyme activity, and reinforcement of root&#x2013;soil interactions through exopolysaccharide production and root surface colonization. In salt-affected paddy systems, inoculated rice plants exhibited reduced sodium uptake, enhanced osmolyte accumulation, and improved nutrient use efficiency, allowing biomass yields to be maintained without synthetic inputs (<xref ref-type="bibr" rid="ref77">Marzouk et al., 2025</xref>; <xref ref-type="bibr" rid="ref119">Sackey et al., 2025</xref>; <xref ref-type="bibr" rid="ref120">Saleem et al., 2025</xref>). However, the performance of these consortia remains highly variable. Inoculant efficacy depends on cultivar compatibility, soil physicochemical properties, and previous land-use history, which collectively shape the recruitment, colonization, and functional integration of applied microbes. However, most studies have focused on short-term growth and yield outcomes, paying limited attention to long-term microbiome restructuring, ecological trade-offs, or economic scalability.</p>
</sec>
</sec>
<sec id="sec23">
<label>7</label>
<title>Challenges and limitations of microbial BCAs</title>
<p>Despite the expanding role of microbial BCAs in sustainable agriculture, their inconsistent performance under field conditions remains a major problem. Laboratory and greenhouse trials often yield promising results, but efficacy frequently declines in open-field systems due to the combined effects of fluctuating soil physicochemical properties, climatic variability, and the ecological complexity of indigenous microbiota. Context-dependent interactions between the inoculant, host genotype, and native microbial networks can either facilitate or hinder colonization, while abiotic stresses such as drought and salinity further compromise microbial viability and plant&#x2013;microbe signalling.</p>
<p>An important challenge is the short-lived colonization of many BCAs, including widely used agents such as <italic>Trichoderma</italic> and <italic>Bacillus</italic> spp. Competitive exclusion by resident microbiota, predation by protists, and suboptimal environmental cues often prevent long-term persistence. Under such conditions, many protective traits, including mycoparasitism, antibiosis, and ISR are regulated by density-dependent microbial signalling, secondary metabolite production, and host-derived signalling cascades. Consequently, incomplete establishment can render BCAs functionally ineffective. While multi-strain consortia and synthetic microbial communities (SynComs) offer a promising avenue to improve stability and broaden functional capacity, mechanistic understanding of interspecies interactions remains limited. Synergistic combinations can enhance disease suppression and stress tolerance, yet antagonistic effects or suppression by native soil communities are also possible. Moreover, the interplay between consortium performance and abiotic stress adaptation is poorly characterized, constraining the development of climate-resilient inoculants.</p>
<p>Regulatory and adoption barriers further limit BCA deployment. Registration frameworks are often fragmented and slow to adapt to living, strain-specific products, especially for consortia or engineered microbes. In many regions, inconsistent biosafety evaluation, quality control, and shelf-life standards undermine both market entry and farmer confidence. Adoption is further constrained by limited farmer awareness, inadequate technical support, and the perception that BCAs are less reliable than synthetic pesticides under high disease pressure or adverse weather.</p>
</sec>
<sec id="sec24">
<label>8</label>
<title>Conclusions and future directions</title>
<p>As agriculture transitions toward sustainable and climate-resilient production systems, microbial BCAs and rhizosphere microbiome-based interventions are expected to become integral components of crop protection and productivity frameworks. Realizing this potential will require a shift from unsystematic inoculation practices toward precision-designed, ecologically compatible, and context-specific microbial solutions. Rationally assembled SynComs offer significant advantages over conventional single-strain inoculants through the integration of complementary functions, including nitrogen fixation, phosphate solubilization, siderophore production, and stress-induced priming. This functional integration also confers redundancy and ecological stability under variable field conditions. Advances in high-throughput omics, including metagenomics, transcriptomics, and metabolomics, now enable the identification of microbial candidates based on functional traits, stress-resilience genes, and biosynthetic potential, rather than cultivability alone. Coupled with genome editing and synthetic biology, these approaches could accelerate the development of functionally optimized BCAs, tailored to specific soil types, climatic regimes, and crop genotypes. However, approaches relying on synthetic biology, genome editing, and fully engineered SynComs largely represent longer-term opportunities, as their field validation, regulatory approval, and ecological risk assessment remain ongoing challenges.</p>
<p>Integration of BCAs into microbiome-informed IPM frameworks will be essential to ensure both disease suppression and long-term ecosystem functionality. Microbiome-informed biological control emphasizes the selection and deployment of BCAs based on key functional traits including stress tolerance, niche complementarity, persistence under field conditions, and the capacity to interact synergistically with host plants and resident microbiomes. In the near term, strategies such as trait-based strain selection, formulation optimization, and the integration of BCAs into existing IPM programs offer more immediately feasible pathways for improving disease management and stress resilience. Emerging precision agriculture tools such as real-time microbiome diagnostics, drone-assisted delivery, and GPS-guided application can enable site-specific, optimized deployment that maximizes efficacy while minimizing non-target impacts. Equally, leveraging the plant holobiome concept in breeding programs offers a path toward host genotypes inherently predisposed to recruit and sustain beneficial microbiota under stress. Embedding microbiome responsiveness into genetic selection criteria, could improve future crops&#x2019; colonization efficiency and functional engagement with BCAs, fostering durable resilience in dynamic agroecosystems. Another challenge is that despite extensive research, quantitative comparisons of BCAs&#x2019; efficacy across crops, environments, and stress conditions remain limited, reflecting heterogeneity in study designs and response metrics as well as the scarcity of standardized field validation.</p>
<p>Ultimately, sustaining yields and ecological integrity under accelerating climatic and biotic pressures will require adaptive, multifunctional microbial solutions developed through a convergence of microbiome science, plant genetics, ecological modelling, and precision delivery technologies. In this context, I propose a conceptual framework (<xref ref-type="fig" rid="fig1">Figure 1</xref>) that advances existing IPM and microbiome-engineering paradigms by integrating (i) microbial design and synthetic consortia, (ii) omics-based screening and functional optimization, (iii) microbiome-informed IPM and precision agriculture, and (iv) host genetics and holobiont optimization across environmental gradients. Through this broad scope, the framework explicitly links near-term, deployable interventions to longer-term innovation pathways, moving beyond descriptive biocontrol strategies toward a systems-level approach for resilient, climate-smart agriculture.</p>
</sec>
</body>
<back>
<sec sec-type="author-contributions" id="sec25">
<title>Author contributions</title>
<p>MI: Conceptualization, Data curation, Funding acquisition, Investigation, Methodology, Project administration, Resources, Writing &#x2013; original draft, Writing &#x2013; review &#x0026; editing.</p>
</sec>
<ack>
<title>Acknowledgments</title>
<p>I sincerely thank Johan A. Stenberg for his insightful comments and critical input during the preparation of this review. I also gratefully acknowledge the handling editor, AA, and the reviewers for their constructive feedback, valuable suggestions, and time, which substantially improved the quality of the manuscript.</p>
</ack>
<sec sec-type="COI-statement" id="sec26">
<title>Conflict of interest</title>
<p>The author(s) declared that this work was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec sec-type="ai-statement" id="sec27">
<title>Generative AI statement</title>
<p>The author(s) declared that Generative AI was not used in the creation of this manuscript.</p>
<p>Any alternative text (alt text) provided alongside figures in this article has been generated by Frontiers with the support of artificial intelligence and reasonable efforts have been made to ensure accuracy, including review by the authors wherever possible. If you identify any issues, please contact us.</p>
</sec>
<sec sec-type="disclaimer" id="sec28">
<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>
<ref-list>
<title>References</title>
<ref id="ref1"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Abdalla</surname><given-names>M.</given-names></name> <name><surname>Bitterlich</surname><given-names>M.</given-names></name> <name><surname>Jansa</surname><given-names>J.</given-names></name> <name><surname>P&#x00FC;schel</surname><given-names>D.</given-names></name> <name><surname>Ahmed</surname><given-names>M. A.</given-names></name></person-group> (<year>2023</year>). <article-title>The role of arbuscular mycorrhizal symbiosis in improving plant water status under drought</article-title>. <source>J. Exp. Bot.</source> <volume>74</volume>, <fpage>4808</fpage>&#x2013;<lpage>4824</lpage>. doi: <pub-id pub-id-type="doi">10.1093/jxb/erad249</pub-id>, <pub-id pub-id-type="pmid">37409696</pub-id></mixed-citation></ref>
<ref id="ref2"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Ali</surname><given-names>D. F. I.</given-names></name> <name><surname>El-Nahrawy</surname><given-names>S.</given-names></name> <name><surname>El-Zawawy</surname><given-names>H. A.</given-names></name> <name><surname>Omara</surname><given-names>A. E. D.</given-names></name></person-group> (<year>2025</year>). <article-title>Effective applications of <italic>Bacillus subtilis</italic> and <italic>B. amyloliquefaciens</italic> as biocontrol agents of damping-off disease and biostimulation of tomato plants</article-title>. <source>Stress</source> <volume>5</volume>:<fpage>9</fpage>. doi: <pub-id pub-id-type="doi">10.3390/stresses5010009</pub-id></mixed-citation></ref>
<ref id="ref3"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Altomare</surname><given-names>C.</given-names></name> <name><surname>Norvell</surname><given-names>W. A.</given-names></name> <name><surname>Bj&#x00F6;rkman</surname><given-names>T.</given-names></name> <name><surname>Harman</surname><given-names>G.</given-names></name></person-group> (<year>1999</year>). <article-title>Solubilization of phosphates and micronutrients by the plant-growth-promoting and biocontrol fungus <italic>Trichoderma harzianum</italic> Rifai 1295-22</article-title>. <source>Appl. Environ. Microbiol.</source> <volume>65</volume>, <fpage>2926</fpage>&#x2013;<lpage>2933</lpage>.</mixed-citation></ref>
<ref id="ref4"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Al-Turki</surname><given-names>A.</given-names></name> <name><surname>Murali</surname><given-names>M.</given-names></name> <name><surname>Omar</surname><given-names>A. F.</given-names></name> <name><surname>Rehan</surname><given-names>M.</given-names></name> <name><surname>Sayyed</surname><given-names>R. Z.</given-names></name></person-group> (<year>2023</year>). <article-title>Recent advances in PGPR-mediated resilience toward interactive effects of drought and salt stress in plants</article-title>. <source>Front. Microbiol.</source> <volume>14</volume>:<fpage>1214845</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fmicb.2023.1214845</pub-id>, <pub-id pub-id-type="pmid">37829451</pub-id></mixed-citation></ref>
<ref id="ref5"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Andronov</surname><given-names>E. E.</given-names></name> <name><surname>Petrova</surname><given-names>S. N.</given-names></name> <name><surname>Pinaev</surname><given-names>A. G.</given-names></name> <name><surname>Pershina</surname><given-names>E. V.</given-names></name> <name><surname>Rakhimgalieva</surname><given-names>S. Z.</given-names></name> <name><surname>Akhmedenov</surname><given-names>K. M.</given-names></name> <etal/></person-group>. (<year>2012</year>). <article-title>Analysis of the structure of microbial community in soils with different degrees of salinization using T-RFLP and real-time PCR techniques</article-title>. <source>Eurasian Soil Sci.</source> <volume>45</volume>, <fpage>147</fpage>&#x2013;<lpage>156</lpage>. doi: <pub-id pub-id-type="doi">10.1134/s1064229312020044</pub-id></mixed-citation></ref>
<ref id="ref6"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Ansabayeva</surname><given-names>A.</given-names></name> <name><surname>Makhambetov</surname><given-names>M.</given-names></name> <name><surname>Rebouh</surname><given-names>N. Y.</given-names></name> <name><surname>Abdelkader</surname><given-names>M.</given-names></name> <name><surname>Saudy</surname><given-names>H. S.</given-names></name> <name><surname>Hassan</surname><given-names>K. M.</given-names></name> <etal/></person-group>. (<year>2025</year>). <article-title>Plant growth-promoting microbes for resilient farming systems: mitigating environmental stressors and boosting crops productivity&#x2014;a review</article-title>. <source>Horticulture</source> <volume>11</volume>:<fpage>260</fpage>. doi: <pub-id pub-id-type="doi">10.3390/horticulturae11030260</pub-id></mixed-citation></ref>
<ref id="ref7"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Asghar</surname><given-names>U.</given-names></name> <name><surname>Malik</surname><given-names>M. F.</given-names></name> <name><surname>Javed</surname><given-names>A.</given-names></name></person-group> (<year>2016</year>). <article-title>Pesticide exposure and human health: a review</article-title>. <source>J. Ecosys. Ecograph.</source> <volume>S5</volume>:<fpage>5</fpage>. doi: <pub-id pub-id-type="doi">10.4172/2157-7625.S5-005</pub-id></mixed-citation></ref>
<ref id="ref8"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Balla</surname><given-names>A.</given-names></name> <name><surname>Silini</surname><given-names>A.</given-names></name> <name><surname>Cherif-Silini</surname><given-names>H.</given-names></name> <name><surname>Chenari Bouket</surname><given-names>A.</given-names></name> <name><surname>Alenezi</surname><given-names>F. N.</given-names></name> <name><surname>Belbahri</surname><given-names>L.</given-names></name></person-group> (<year>2022</year>). <article-title>Recent advances in encapsulation techniques of plant growth-promoting microorganisms and their prospects in the sustainable agriculture</article-title>. <source>Appl. Sci.</source> <volume>12</volume>:<fpage>9020</fpage>. doi: <pub-id pub-id-type="doi">10.3390/app12189020</pub-id></mixed-citation></ref>
<ref id="ref9"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Bebber</surname><given-names>D. P.</given-names></name> <name><surname>Ramotowski</surname><given-names>M. A.</given-names></name> <name><surname>Gurr</surname><given-names>S. J.</given-names></name></person-group> (<year>2013</year>). <article-title>Crop pests and pathogens move polewards in a warming world</article-title>. <source>Nat. Clim. Chang.</source> <volume>3</volume>, <fpage>985</fpage>&#x2013;<lpage>988</lpage>. doi: <pub-id pub-id-type="doi">10.1038/nclimate1990</pub-id></mixed-citation></ref>
<ref id="ref10"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Berendsen</surname><given-names>R. L.</given-names></name> <name><surname>Pieterse</surname><given-names>C. M.</given-names></name> <name><surname>Bakker</surname><given-names>P. A.</given-names></name></person-group> (<year>2012</year>). <article-title>The rhizosphere microbiome and plant health</article-title>. <source>Trends Plant Sci.</source> <volume>17</volume>, <fpage>478</fpage>&#x2013;<lpage>486</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.tplants.2012.04.001</pub-id>, <pub-id pub-id-type="pmid">22564542</pub-id></mixed-citation></ref>
<ref id="ref11"><mixed-citation publication-type="book"><person-group person-group-type="author"><name><surname>Berg</surname><given-names>G.</given-names></name> <name><surname>Alavi</surname><given-names>M.</given-names></name> <name><surname>Schmidt</surname><given-names>C. S.</given-names></name> <name><surname>Zachow</surname><given-names>C.</given-names></name> <name><surname>Egamberdieva</surname><given-names>D.</given-names></name> <name><surname>Kamilova</surname><given-names>F.</given-names></name> <etal/></person-group>. (<year>2013</year>). &#x201C;<chapter-title>Biocontrol and osmoprotection for plants under saline conditions</chapter-title>&#x201D; in <source>Molecular Microbial Ecology of the Rhizosphere</source>. ed. <person-group person-group-type="editor"><name><surname>de Bruijn</surname><given-names>F. J.</given-names></name></person-group>. (<publisher-loc>Hoboken, NJ</publisher-loc>: <publisher-name>Wiley-Blackwell</publisher-name>).</mixed-citation></ref>
<ref id="ref12"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Berg</surname><given-names>G.</given-names></name> <name><surname>Smalla</surname><given-names>K.</given-names></name></person-group> (<year>2009</year>). <article-title>Plant species and soil type cooperatively shape the structure and function of microbial communities in the rhizosphere</article-title>. <source>FEMS Microbiol. Ecol.</source> <volume>68</volume>, <fpage>1</fpage>&#x2013;<lpage>13</lpage>. doi: <pub-id pub-id-type="doi">10.1111/j.1574-6941.2009.00654.x</pub-id>, <pub-id pub-id-type="pmid">19243436</pub-id></mixed-citation></ref>
<ref id="ref13"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Busby</surname><given-names>P. E.</given-names></name> <name><surname>Soman</surname><given-names>C.</given-names></name> <name><surname>Wagner</surname><given-names>M. R.</given-names></name> <name><surname>Friesen</surname><given-names>M. L.</given-names></name> <name><surname>Kremer</surname><given-names>J.</given-names></name> <name><surname>Bennett</surname><given-names>A.</given-names></name> <etal/></person-group>. (<year>2017</year>). <article-title>Research priorities for harnessing plant microbiomes in sustainable agriculture</article-title>. <source>PLoS Biol.</source> <volume>15</volume>:<fpage>e2001793</fpage>. doi: <pub-id pub-id-type="doi">10.1371/journal.pbio.2001793</pub-id>, <pub-id pub-id-type="pmid">28350798</pub-id></mixed-citation></ref>
<ref id="ref14"><mixed-citation publication-type="book"><person-group person-group-type="author"><name><surname>Chandel</surname><given-names>A. K.</given-names></name> <name><surname>Chen</surname><given-names>H.</given-names></name> <name><surname>Sharma</surname><given-names>H. C.</given-names></name> <name><surname>Adhikari</surname><given-names>K.</given-names></name> <name><surname>Gao</surname><given-names>B.</given-names></name></person-group> (<year>2019</year>). &#x201C;<chapter-title>Beneficial microbes for sustainable agriculture</chapter-title>&#x201D; in eds. R. Chandra and RC Sobti. <source>Microbes for sustainable development and bioremediation</source>. (<publisher-loc>Boca Raton</publisher-loc>: <publisher-name>CRC press</publisher-name>), <fpage>257</fpage>&#x2013;<lpage>265</lpage>.</mixed-citation></ref>
<ref id="ref15"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Chapelle</surname><given-names>E.</given-names></name> <name><surname>Mendes</surname><given-names>R.</given-names></name> <name><surname>Bakker</surname><given-names>P. A.</given-names></name> <name><surname>Raaijmakers</surname><given-names>J. M.</given-names></name></person-group> (<year>2016</year>). <article-title>Fungal invasion of the rhizosphere microbiome</article-title>. <source>ISME J.</source> <volume>10</volume>, <fpage>265</fpage>&#x2013;<lpage>268</lpage>. doi: <pub-id pub-id-type="doi">10.1038/ismej.2015.82</pub-id></mixed-citation></ref>
<ref id="ref16"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Chen</surname><given-names>Q.</given-names></name> <name><surname>Song</surname><given-names>Y.</given-names></name> <name><surname>An</surname><given-names>Y.</given-names></name> <name><surname>Lu</surname><given-names>Y.</given-names></name> <name><surname>Zhong</surname><given-names>G.</given-names></name></person-group> (<year>2024</year>). <article-title>Mechanisms and impact of rhizosphere microbial metabolites on crop health, traits, functional components: a comprehensive review</article-title>. <source>Molecules</source> <volume>29</volume>:<fpage>5922</fpage>. doi: <pub-id pub-id-type="doi">10.3390/molecules29245922</pub-id></mixed-citation></ref>
<ref id="ref17"><mixed-citation publication-type="book"><person-group person-group-type="author"><name><surname>Chet</surname><given-names>I.</given-names></name></person-group> ed. (<year>1987</year>). &#x201C;<chapter-title>Trichoderma: application, mode of action, and potential as biocontrol agent of soilborne plant pathogenic fungi</chapter-title>&#x201D; in <source>Innovative approaches to Plant Disease Control</source>. (<publisher-loc>New York</publisher-loc>: <publisher-name>John Wiley</publisher-name>), <fpage>137</fpage>&#x2013;<lpage>160</lpage>.</mixed-citation></ref>
<ref id="ref18"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Chevallereau</surname><given-names>A.</given-names></name> <name><surname>Pons</surname><given-names>B. J.</given-names></name> <name><surname>van Houte</surname><given-names>S.</given-names></name> <name><surname>Westra</surname><given-names>E. R.</given-names></name></person-group> (<year>2022</year>). <article-title>Interactions between bacterial and phage communities in natural environments</article-title>. <source>Nat. Rev. Microbiol.</source> <volume>20</volume>, <fpage>49</fpage>&#x2013;<lpage>62</lpage>. doi: <pub-id pub-id-type="doi">10.1038/s41579-021-00602-y</pub-id></mixed-citation></ref>
<ref id="ref19"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Choudhary</surname><given-names>A.</given-names></name> <name><surname>Senthil-Kumar</surname><given-names>M.</given-names></name></person-group> (<year>2024</year>). <article-title>Drought: a context-dependent damper and aggravator of plant diseases</article-title>. <source>Plant Cell Environ.</source> <volume>47</volume>, <fpage>2109</fpage>&#x2013;<lpage>2126</lpage>. doi: <pub-id pub-id-type="doi">10.1111/pce.14863</pub-id></mixed-citation></ref>
<ref id="ref20"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Chukwuneme</surname><given-names>C. F.</given-names></name> <name><surname>Babalola</surname><given-names>O. O.</given-names></name></person-group> (<year>2025</year>). <article-title>Microbial diversity and function in the rhizosphere microbiome: driving forces and monitoring approaches</article-title>. <source>Agrosyst. Geosci. Environ.</source> <volume>8</volume>:<fpage>e70169</fpage>. doi: <pub-id pub-id-type="doi">10.1002/agg2.70169</pub-id></mixed-citation></ref>
<ref id="ref21"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Compant</surname><given-names>S.</given-names></name> <name><surname>Samad</surname><given-names>A.</given-names></name> <name><surname>Faist</surname><given-names>H.</given-names></name> <name><surname>Sessitsch</surname><given-names>A.</given-names></name></person-group> (<year>2019</year>). <article-title>A review on the plant microbiome: ecology, functions, and emerging trends in microbial application</article-title>. <source>J. Adv. Res.</source> <volume>19</volume>, <fpage>29</fpage>&#x2013;<lpage>37</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.jare.2019.03.004</pub-id>, <pub-id pub-id-type="pmid">31341667</pub-id></mixed-citation></ref>
<ref id="ref22"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Cruz-Barrera</surname><given-names>M.</given-names></name> <name><surname>Izquierdo-Garc&#x00ED;a</surname><given-names>L. F.</given-names></name> <name><surname>G&#x00F3;mez-Marroqu&#x00ED;n</surname><given-names>M.</given-names></name> <name><surname>Santos-D&#x00ED;az</surname><given-names>A.</given-names></name> <name><surname>Uribe-Guti&#x00E9;rrez</surname><given-names>L.</given-names></name> <name><surname>Moreno-Velandia</surname><given-names>C. A.</given-names></name></person-group> (<year>2024</year>). <article-title>Hydrogel capsules as new delivery system for <italic>Trichoderma koningiopsis</italic> Th003 to control <italic>Rhizoctonia solani</italic> in rice (<italic>Oryza sativa</italic>)</article-title>. <source>World J. Microbiol. Biotechnol.</source> <volume>40</volume>:<fpage>108</fpage>. doi: <pub-id pub-id-type="doi">10.1007/s11274-024-03897-0</pub-id></mixed-citation></ref>
<ref id="ref23"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Damodaran</surname><given-names>T.</given-names></name> <name><surname>Rajan</surname><given-names>S.</given-names></name> <name><surname>Muthukumar</surname><given-names>M.</given-names></name> <name><surname>Gopal</surname><given-names>R.</given-names></name> <name><surname>Yadav</surname><given-names>K.</given-names></name> <name><surname>Kumar</surname><given-names>S.</given-names></name> <etal/></person-group>. (<year>2020</year>). <article-title>Biological management of banana fusarium wilt caused by <italic>fusarium oxysporum</italic> f. sp. <italic>cubense</italic> tropical race 4 using antagonistic fungal isolate CSR-T-3 (<italic>Trichoderma reesei</italic>)</article-title>. <source>Front. Microbiol.</source> <volume>11</volume>:<fpage>595845</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fmicb.2020.595845</pub-id></mixed-citation></ref>
<ref id="ref24"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>De Faria</surname><given-names>M. R.</given-names></name> <name><surname>Costa</surname><given-names>L. S. A. S.</given-names></name> <name><surname>Chiaramonte</surname><given-names>J. B.</given-names></name> <name><surname>Bettiol</surname><given-names>W.</given-names></name> <name><surname>Mendes</surname><given-names>R.</given-names></name></person-group> (<year>2021</year>). <article-title>The rhizosphere microbiome: functions, dynamics, and role in plant protection</article-title>. <source>Trop. Plant Pathol.</source> <volume>46</volume>, <fpage>13</fpage>&#x2013;<lpage>25</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s40858-020-00390-5</pub-id></mixed-citation></ref>
<ref id="ref25"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>De Vries</surname><given-names>F. T.</given-names></name> <name><surname>Wallenstein</surname><given-names>M. D.</given-names></name></person-group> (<year>2017</year>). <article-title>Below-ground connections underlying above-ground food production: a framework for optimising ecological connections in the rhizosphere</article-title>. <source>J. Ecol.</source> <volume>105</volume>, <fpage>913</fpage>&#x2013;<lpage>920</lpage>. doi: <pub-id pub-id-type="doi">10.1111/1365-2745.12783</pub-id></mixed-citation></ref>
<ref id="ref26"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Debode</surname><given-names>J.</given-names></name> <name><surname>De Tender</surname><given-names>C.</given-names></name> <name><surname>Cremelie</surname><given-names>P.</given-names></name> <name><surname>Lee</surname><given-names>A. S.</given-names></name> <name><surname>Kyndt</surname><given-names>T.</given-names></name> <name><surname>Muylle</surname><given-names>H.</given-names></name> <etal/></person-group>. (<year>2018</year>). <article-title><italic>Trichoderma</italic>-inoculated miscanthus straw can replace peat in strawberry cultivation, with beneficial effects on disease control</article-title>. <source>Front. Plant Sci.</source> <volume>9</volume>:<fpage>213</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fpls.2018.00213</pub-id></mixed-citation></ref>
<ref id="ref27"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Deng</surname><given-names>X.</given-names></name> <name><surname>Zhang</surname><given-names>N.</given-names></name> <name><surname>Shen</surname><given-names>Z.</given-names></name> <name><surname>Zhu</surname><given-names>C.</given-names></name> <name><surname>Liu</surname><given-names>H.</given-names></name> <name><surname>Xu</surname><given-names>Z.</given-names></name> <etal/></person-group>. (<year>2021</year>). <article-title>Soil microbiome manipulation triggers direct and possible indirect suppression against <italic>Ralstonia solanacearum</italic> and <italic>fusarium oxysporum</italic></article-title>. <source>NPJ Biofilms Microbiomes</source> <volume>7</volume>:<fpage>33</fpage>. doi: <pub-id pub-id-type="doi">10.1038/s41522-021-00204-9</pub-id></mixed-citation></ref>
<ref id="ref28"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Devi</surname><given-names>Y. B.</given-names></name> <name><surname>Meetei</surname><given-names>T. T.</given-names></name> <name><surname>Kumari</surname><given-names>N.</given-names></name></person-group> (<year>2018</year>). <article-title>Impact of pesticides on soil microbial diversity and enzymes: a review</article-title>. <source>Int. J. Curr. Microbiol. Appl. Sci.</source> <volume>7</volume>, <fpage>952</fpage>&#x2013;<lpage>958</lpage>. doi: <pub-id pub-id-type="doi">10.20546/ijcmas.2018.706.113</pub-id></mixed-citation></ref>
<ref id="ref29"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Di Marco</surname><given-names>S.</given-names></name> <name><surname>Osti</surname><given-names>F.</given-names></name> <name><surname>Cesari</surname><given-names>A.</given-names></name></person-group> (<year>2004</year>). <article-title>Experiments on the control of esca by <italic>Trichoderma</italic></article-title>. <source>Phytopathol. Mediterr.</source> <volume>43</volume>, <fpage>108</fpage>&#x2013;<lpage>115</lpage>. doi: <pub-id pub-id-type="doi">10.36253/phyto-5040</pub-id></mixed-citation></ref>
<ref id="ref30"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Doni</surname><given-names>F.</given-names></name> <name><surname>Fathurrahman</surname><given-names>F.</given-names></name> <name><surname>Mispan</surname><given-names>M. S.</given-names></name> <name><surname>Suhaimi</surname><given-names>N. S. M.</given-names></name> <name><surname>Yusoff</surname><given-names>W. M. W.</given-names></name> <name><surname>Uphoff</surname><given-names>N.</given-names></name></person-group> (<year>2019</year>). <article-title>Transcriptomic profiling of rice seedlings inoculated with the symbiotic fungus <italic>Trichoderma asperellum</italic> SL2</article-title>. <source>J. Plant Growth Regul.</source> <volume>38</volume>, <fpage>1507</fpage>&#x2013;<lpage>1515</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s00344-019-09952-7</pub-id></mixed-citation></ref>
<ref id="ref31"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Fierer</surname><given-names>N.</given-names></name></person-group> (<year>2017</year>). <article-title>Embracing the unknown: disentangling the complexities of the soil microbiome</article-title>. <source>Nat. Rev. Microbiol.</source> <volume>15</volume>, <fpage>579</fpage>&#x2013;<lpage>590</lpage>. doi: <pub-id pub-id-type="doi">10.1038/nrmicro.2017.87</pub-id>, <pub-id pub-id-type="pmid">28824177</pub-id></mixed-citation></ref>
<ref id="ref32"><mixed-citation publication-type="book"><person-group person-group-type="author"><name><surname>Fischer</surname><given-names>S.</given-names></name> <name><surname>Pr&#x00ED;ncipe</surname><given-names>A.</given-names></name> <name><surname>Alvarez</surname><given-names>F.</given-names></name> <name><surname>Cordero</surname><given-names>P.</given-names></name> <name><surname>Castro</surname><given-names>M.</given-names></name> <name><surname>Godino</surname><given-names>A.</given-names></name> <etal/></person-group>. (<year>2013</year>). &#x201C;<chapter-title>Fighting plant diseases through the application of <italic>Bacillus</italic> and <italic>Pseudomonas</italic> strains</chapter-title>&#x201D; in ed. R. Aroca. <source>Symbiotic Endophytes</source> (<publisher-loc>Berlin, Heidelberg</publisher-loc>: <publisher-name>Springer</publisher-name>), <fpage>165</fpage>&#x2013;<lpage>193</lpage>.</mixed-citation></ref>
<ref id="ref33"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Fisher</surname><given-names>M. C.</given-names></name> <name><surname>Hawkins</surname><given-names>N. J.</given-names></name> <name><surname>Sanglard</surname><given-names>D.</given-names></name> <name><surname>Gurr</surname><given-names>S. J.</given-names></name></person-group> (<year>2018</year>). <article-title>Worldwide emergence of resistance to antifungal drugs challenges human health and food security</article-title>. <source>Science</source> <volume>360</volume>, <fpage>739</fpage>&#x2013;<lpage>742</lpage>. doi: <pub-id pub-id-type="doi">10.1126/science.aap7999</pub-id>, <pub-id pub-id-type="pmid">29773744</pub-id></mixed-citation></ref>
<ref id="ref34"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Freeman</surname><given-names>S.</given-names></name> <name><surname>Minz</surname><given-names>D.</given-names></name> <name><surname>Kolesnik</surname><given-names>I.</given-names></name> <name><surname>Barbul</surname><given-names>O.</given-names></name> <name><surname>Zveibil</surname><given-names>A.</given-names></name> <name><surname>Maymon</surname><given-names>M.</given-names></name> <etal/></person-group>. (<year>2004</year>). <article-title><italic>Trichoderma</italic> biocontrol of <italic>Colletotrichum acutatum</italic> and <italic>Botrytis cinerea</italic> and survival in strawberry</article-title>. <source>Eur. J. Plant Pathol.</source> <volume>110</volume>, <fpage>361</fpage>&#x2013;<lpage>370</lpage>. doi: <pub-id pub-id-type="doi">10.1023/b:ejpp.0000021057.93305.d9</pub-id></mixed-citation></ref>
<ref id="ref35"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Gao</surname><given-names>P.</given-names></name> <name><surname>Qi</surname><given-names>K.</given-names></name> <name><surname>Han</surname><given-names>Y.</given-names></name> <name><surname>Ma</surname><given-names>L.</given-names></name> <name><surname>Zhang</surname><given-names>B.</given-names></name> <name><surname>Zhang</surname><given-names>Y.</given-names></name> <etal/></person-group>. (<year>2023</year>). <article-title>Effect of <italic>Trichoderma viride</italic> on rhizosphere microbial communities and biocontrol of soybean root rot</article-title>. <source>Front. Microbiol.</source> <volume>14</volume>:<fpage>1204688</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fmicb.2023.1204688</pub-id>, <pub-id pub-id-type="pmid">37333630</pub-id></mixed-citation></ref>
<ref id="ref36"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Geng</surname><given-names>Y.</given-names></name> <name><surname>Chen</surname><given-names>S.</given-names></name> <name><surname>Lv</surname><given-names>P.</given-names></name> <name><surname>Li</surname><given-names>Y.</given-names></name> <name><surname>Li</surname><given-names>J.</given-names></name> <name><surname>Jiang</surname><given-names>F.</given-names></name> <etal/></person-group>. (<year>2025</year>). <article-title>Positive role of <italic>Trichoderma harzianum</italic> in increasing plant tolerance to abiotic stresses: a review</article-title>. <source>Antioxidants</source> <volume>14</volume>:<fpage>807</fpage>. doi: <pub-id pub-id-type="doi">10.3390/antiox14070807</pub-id>, <pub-id pub-id-type="pmid">40722911</pub-id></mixed-citation></ref>
<ref id="ref37"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Goulson</surname><given-names>D.</given-names></name> <name><surname>Nicholls</surname><given-names>E.</given-names></name> <name><surname>Bot&#x00ED;as</surname><given-names>C.</given-names></name> <name><surname>Rotheray</surname><given-names>E. L.</given-names></name></person-group> (<year>2015</year>). <article-title>Bee declines driven by combined stress from parasites, pesticides, and lack of flowers</article-title>. <source>Science</source> <volume>347</volume>:<fpage>1255957</fpage>. doi: <pub-id pub-id-type="doi">10.1126/science.1255957</pub-id>, <pub-id pub-id-type="pmid">25721506</pub-id></mixed-citation></ref>
<ref id="ref38"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Gramaje</surname><given-names>D.</given-names></name> <name><surname>Urbez-Torres</surname><given-names>J. R.</given-names></name> <name><surname>Sosnowski</surname><given-names>M. R.</given-names></name></person-group> (<year>2018</year>). <article-title>Managing grapevine trunk diseases with respect to etiology and epidemiology: current strategies and future prospects</article-title>. <source>Plant Dis.</source> <volume>102</volume>, <fpage>12</fpage>&#x2013;<lpage>39</lpage>. doi: <pub-id pub-id-type="doi">10.1094/PDIS-04-17-0512-FE</pub-id>, <pub-id pub-id-type="pmid">30673457</pub-id></mixed-citation></ref>
<ref id="ref39"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Gu</surname><given-names>Y.</given-names></name> <name><surname>Liang</surname><given-names>W.</given-names></name> <name><surname>Li</surname><given-names>Z.</given-names></name> <name><surname>Liu</surname><given-names>S.</given-names></name> <name><surname>Liang</surname><given-names>S.</given-names></name> <name><surname>Lei</surname><given-names>P.</given-names></name> <etal/></person-group>. (<year>2023</year>). <article-title>The biocontrol agent <italic>Bacillus velezensis</italic> T-5 changes the soil bacterial community composition by affecting the tomato root exudate profile</article-title>. <source>Plant Soil</source> <volume>490</volume>, <fpage>669</fpage>&#x2013;<lpage>680</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s11104-023-06114-3</pub-id></mixed-citation></ref>
<ref id="ref40"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Guzm&#x00E1;n-Guzm&#x00E1;n</surname><given-names>P.</given-names></name> <name><surname>Porras-Troncoso</surname><given-names>M. D.</given-names></name> <name><surname>Olmedo-Monfil</surname><given-names>V.</given-names></name> <name><surname>Herrera-Estrella</surname><given-names>A.</given-names></name></person-group> (<year>2019</year>). <article-title><italic>Trichoderma</italic> species: versatile plant symbionts</article-title>. <source>Phytopathology</source> <volume>109</volume>, <fpage>6</fpage>&#x2013;<lpage>16</lpage>. doi: <pub-id pub-id-type="doi">10.1094/PHYTO-07-18-0218-RVW</pub-id>, <pub-id pub-id-type="pmid">30412012</pub-id></mixed-citation></ref>
<ref id="ref41"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Haichar</surname><given-names>F. E. Z.</given-names></name> <name><surname>Marol</surname><given-names>C.</given-names></name> <name><surname>Berge</surname><given-names>O.</given-names></name> <name><surname>Rangel-Castro</surname><given-names>J. I.</given-names></name> <name><surname>Prosser</surname><given-names>J. I.</given-names></name> <name><surname>Balesdent</surname><given-names>J.</given-names></name> <etal/></person-group>. (<year>2008</year>). <article-title>Plant host habitat and root exudates shape soil bacterial community structure</article-title>. <source>ISME J.</source> <volume>2</volume>, <fpage>1221</fpage>&#x2013;<lpage>1230</lpage>. doi: <pub-id pub-id-type="doi">10.1038/ismej.2008.80</pub-id>, <pub-id pub-id-type="pmid">18754043</pub-id></mixed-citation></ref>
<ref id="ref42"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Han</surname><given-names>S. W.</given-names></name> <name><surname>Yoshikuni</surname><given-names>Y.</given-names></name></person-group> (<year>2022</year>). <article-title>Microbiome engineering for sustainable agriculture: using synthetic biology to enhance nitrogen metabolism in plant-associated microbes</article-title>. <source>Curr. Opin. Microbiol.</source> <volume>68</volume>:<fpage>102172</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.mib.2022.102172</pub-id>, <pub-id pub-id-type="pmid">35717707</pub-id></mixed-citation></ref>
<ref id="ref43"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Ha-Tran</surname><given-names>D. M.</given-names></name> <name><surname>Nguyen</surname><given-names>T. T. M.</given-names></name> <name><surname>Hung</surname><given-names>S. H.</given-names></name> <name><surname>Huang</surname><given-names>E.</given-names></name> <name><surname>Huang</surname><given-names>C. C.</given-names></name></person-group> (<year>2021</year>). <article-title>Roles of plant growth-promoting rhizobacteria (PGPR) in stimulating salinity stress defense in plants: a review</article-title>. <source>Int. J. Mol. Sci.</source> <volume>22</volume>:<fpage>3154</fpage>. doi: <pub-id pub-id-type="doi">10.3390/ijms22063154</pub-id>, <pub-id pub-id-type="pmid">33808829</pub-id></mixed-citation></ref>
<ref id="ref44"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Heil</surname><given-names>M.</given-names></name> <name><surname>Bostock</surname><given-names>R. M.</given-names></name></person-group> (<year>2002</year>). <article-title>Induced systemic resistance (ISR) against pathogens in the context of induced plant defences</article-title>. <source>Ann. Bot.</source> <volume>89</volume>, <fpage>503</fpage>&#x2013;<lpage>512</lpage>. doi: <pub-id pub-id-type="doi">10.1093/aob/mcf076</pub-id></mixed-citation></ref>
<ref id="ref45"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Hermosa</surname><given-names>R.</given-names></name> <name><surname>Rubio</surname><given-names>M. B.</given-names></name> <name><surname>Cardoza</surname><given-names>R. E.</given-names></name> <name><surname>Nicol&#x00E1;s</surname><given-names>C.</given-names></name> <name><surname>Monte</surname><given-names>E.</given-names></name> <name><surname>Guti&#x00E9;rrez</surname><given-names>S.</given-names></name></person-group> (<year>2013</year>). <article-title>The contribution of <italic>Trichoderma</italic> to balancing the costs of plant growth and defense</article-title>. <source>Int. Microbiol.</source> <volume>16</volume>, <fpage>69</fpage>&#x2013;<lpage>80</lpage>. doi: <pub-id pub-id-type="doi">10.2436/20.1501.01.181</pub-id></mixed-citation></ref>
<ref id="ref46"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Hilton</surname><given-names>S.</given-names></name> <name><surname>Bennett</surname><given-names>A. J.</given-names></name> <name><surname>Keane</surname><given-names>G.</given-names></name> <name><surname>Bending</surname><given-names>G. D.</given-names></name> <name><surname>Chandler</surname><given-names>D.</given-names></name> <name><surname>Stobart</surname><given-names>R.</given-names></name> <etal/></person-group>. (<year>2013</year>). <article-title>Impact of shortened crop rotation of oilseed rape on soil and rhizosphere microbial diversity in relation to yield decline</article-title>. <source>PLoS One</source> <volume>8</volume>:<fpage>e59859</fpage>. doi: <pub-id pub-id-type="doi">10.1371/journal.pone.0059859</pub-id>, <pub-id pub-id-type="pmid">23573215</pub-id></mixed-citation></ref>
<ref id="ref47"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Huang</surname><given-names>X.</given-names></name> <name><surname>Zheng</surname><given-names>Y.</given-names></name> <name><surname>Li</surname><given-names>P.</given-names></name> <name><surname>Cui</surname><given-names>J.</given-names></name> <name><surname>Sui</surname><given-names>P.</given-names></name> <name><surname>Chen</surname><given-names>Y.</given-names></name> <etal/></person-group>. (<year>2023</year>). <article-title>Organic management increases beneficial microorganisms and promotes the stability of microecological networks in tea plantation soil</article-title>. <source>Front. Microbiol.</source> <volume>14</volume>:<fpage>1237842</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fmicb.2023.1237842</pub-id>, <pub-id pub-id-type="pmid">37795307</pub-id></mixed-citation></ref>
<ref id="ref48"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Ikeda</surname><given-names>S.</given-names></name> <name><surname>Anda</surname><given-names>M.</given-names></name> <name><surname>Inaba</surname><given-names>S.</given-names></name> <name><surname>Eda</surname><given-names>S.</given-names></name> <name><surname>Sato</surname><given-names>S.</given-names></name> <name><surname>Sasaki</surname><given-names>K.</given-names></name> <etal/></person-group>. (<year>2011</year>). <article-title>Autoregulation of nodulation interferes with impacts of nitrogen fertilization levels on the leaf-associated bacterial community in soybeans</article-title>. <source>Appl. Environ. Microbiol.</source> <volume>77</volume>, <fpage>1973</fpage>&#x2013;<lpage>1980</lpage>. doi: <pub-id pub-id-type="doi">10.1128/AEM.02567-10</pub-id>, <pub-id pub-id-type="pmid">21239540</pub-id></mixed-citation></ref>
<ref id="ref49"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Imran</surname><given-names>A.</given-names></name> <name><surname>Hakim</surname><given-names>S.</given-names></name> <name><surname>Tariq</surname><given-names>M.</given-names></name> <name><surname>Nawaz</surname><given-names>M. S.</given-names></name> <name><surname>Laraib</surname><given-names>I.</given-names></name> <name><surname>Gulzar</surname><given-names>U.</given-names></name> <etal/></person-group>. (<year>2021</year>). <article-title>Diazotrophs for lowering nitrogen pollution crises: looking deep into the roots</article-title>. <source>Front. Microbiol.</source> <volume>12</volume>:<fpage>637815</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fmicb.2021.637815</pub-id>, <pub-id pub-id-type="pmid">34108945</pub-id></mixed-citation></ref>
<ref id="ref50"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Iqbal</surname><given-names>M.</given-names></name> <name><surname>Broberg</surname><given-names>A.</given-names></name> <name><surname>Andreasson</surname><given-names>E.</given-names></name> <name><surname>Stenberg</surname><given-names>J. A.</given-names></name></person-group> (<year>2023</year>). <article-title>Biocontrol potential of beneficial fungus <italic>Aureobasidium pullulans</italic> against <italic>Botrytis cinerea</italic> and <italic>Colletotrichum acutatum</italic></article-title>. <source>Phytopathology</source> <volume>113</volume>, <fpage>1428</fpage>&#x2013;<lpage>1438</lpage>. doi: <pub-id pub-id-type="doi">10.1094/PHYTO-02-23-0067-R</pub-id>, <pub-id pub-id-type="pmid">36945727</pub-id></mixed-citation></ref>
<ref id="ref51"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Iqbal</surname><given-names>M.</given-names></name> <name><surname>Fl&#x00F6;hr</surname><given-names>A.</given-names></name> <name><surname>Andreasson</surname><given-names>E.</given-names></name> <name><surname>Stenberg</surname><given-names>J. A.</given-names></name></person-group> (<year>2025</year>). <article-title>Breeding for integrated pest management (B-IPM): a new concept simultaneously optimising plant resistance and biocontrol</article-title>. <source>Front. Plant Sci.</source> <volume>16</volume>:<fpage>1659069</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fpls.2025.1659069</pub-id>, <pub-id pub-id-type="pmid">41036406</pub-id></mixed-citation></ref>
<ref id="ref52"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Iqbal</surname><given-names>M.</given-names></name> <name><surname>Jamshaid</surname><given-names>M.</given-names></name> <name><surname>Zahid</surname><given-names>M. A.</given-names></name> <name><surname>Andreasson</surname><given-names>E.</given-names></name> <name><surname>Vetukuri</surname><given-names>R. R.</given-names></name> <name><surname>Stenberg</surname><given-names>J. A.</given-names></name></person-group> (<year>2021</year>). <article-title>Biological control of strawberry crown rot, root rot and grey mould by the beneficial fungus <italic>Aureobasidium pullulans</italic></article-title>. <source>BioControl</source> <volume>66</volume>, <fpage>535</fpage>&#x2013;<lpage>545</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s10526-021-10083-w</pub-id></mixed-citation></ref>
<ref id="ref53"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Iqbal</surname><given-names>M.</given-names></name> <name><surname>J&#x00FC;tzeler</surname><given-names>M.</given-names></name> <name><surname>Fran&#x00E7;a</surname><given-names>S. C.</given-names></name> <name><surname>W&#x00E4;ckers</surname><given-names>F.</given-names></name> <name><surname>Andreasson</surname><given-names>E.</given-names></name> <name><surname>Stenberg</surname><given-names>J. A.</given-names></name></person-group> (<year>2022</year>). <article-title>Bee-vectored <italic>Aureobasidium pullulans</italic> for biological control of gray mold in strawberry</article-title>. <source>Phytopathology</source> <volume>112</volume>, <fpage>232</fpage>&#x2013;<lpage>237</lpage>. doi: <pub-id pub-id-type="doi">10.1094/PHYTO-05-21-0205-R</pub-id>, <pub-id pub-id-type="pmid">34181440</pub-id></mixed-citation></ref>
<ref id="ref54"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Jacobsen</surname><given-names>C. S.</given-names></name> <name><surname>Hjelms&#x00F8;</surname><given-names>M. H.</given-names></name></person-group> (<year>2014</year>). <article-title>Agricultural soils, pesticides and microbial diversity</article-title>. <source>Curr. Opin. Biotechnol.</source> <volume>27</volume>, <fpage>15</fpage>&#x2013;<lpage>20</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.copbio.2013.09.003</pub-id>, <pub-id pub-id-type="pmid">24863892</pub-id></mixed-citation></ref>
<ref id="ref55"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Jha</surname><given-names>M.</given-names></name> <name><surname>Chourasia</surname><given-names>S.</given-names></name> <name><surname>Sinha</surname><given-names>S.</given-names></name></person-group> (<year>2013</year>). <article-title>Microbial consortium for sustainable rice production</article-title>. <source>Agroecol. Sustain. Food Syst.</source> <volume>37</volume>, <fpage>340</fpage>&#x2013;<lpage>362</lpage>. doi: <pub-id pub-id-type="doi">10.1080/10440046.2012.672376</pub-id></mixed-citation></ref>
<ref id="ref56"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Kang</surname><given-names>T.</given-names></name> <name><surname>Zhang</surname><given-names>M.</given-names></name> <name><surname>Xia</surname><given-names>M.</given-names></name> <name><surname>Chen</surname><given-names>K.</given-names></name> <name><surname>Zhai</surname><given-names>Y.</given-names></name> <name><surname>Yan</surname><given-names>B.</given-names></name> <etal/></person-group>. (<year>2025</year>). <article-title>Rhizosphere regulation: development and blueprint for soil-borne disease suppression in strawberry</article-title>. <source>Ann. Appl. Biol.</source> <volume>186</volume>, <fpage>27</fpage>&#x2013;<lpage>37</lpage>. doi: <pub-id pub-id-type="doi">10.1111/aab.12961</pub-id></mixed-citation></ref>
<ref id="ref57"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Karlsson</surname><given-names>M.</given-names></name> <name><surname>Durling</surname><given-names>M. B.</given-names></name> <name><surname>Choi</surname><given-names>J.</given-names></name> <name><surname>Kosawang</surname><given-names>C.</given-names></name> <name><surname>Lackner</surname><given-names>G.</given-names></name> <name><surname>Tzelepis</surname><given-names>G. D.</given-names></name> <etal/></person-group>. (<year>2015</year>). <article-title>Insights on the evolution of mycoparasitism from the genome of <italic>Clonostachys rosea</italic></article-title>. <source>Genome Biol. Evol.</source> <volume>7</volume>, <fpage>465</fpage>&#x2013;<lpage>480</lpage>. doi: <pub-id pub-id-type="doi">10.1093/gbe/evu292</pub-id>, <pub-id pub-id-type="pmid">25575496</pub-id></mixed-citation></ref>
<ref id="ref58"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Kashyap</surname><given-names>S.</given-names></name> <name><surname>Sharma</surname><given-names>I.</given-names></name> <name><surname>Dowarah</surname><given-names>B.</given-names></name> <name><surname>Barman</surname><given-names>R.</given-names></name> <name><surname>Gill</surname><given-names>S. S.</given-names></name> <name><surname>Agarwala</surname><given-names>N.</given-names></name></person-group> (<year>2023</year>). <article-title>Plant and soil-associated microbiome dynamics determine the fate of bacterial wilt pathogen <italic>Ralstonia solanacearum</italic></article-title>. <source>Planta</source> <volume>258</volume>:<fpage>57</fpage>. doi: <pub-id pub-id-type="doi">10.1007/s00425-023-04209-w</pub-id>, <pub-id pub-id-type="pmid">37524889</pub-id></mixed-citation></ref>
<ref id="ref59"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Kaur</surname><given-names>R.</given-names></name> <name><surname>Gupta</surname><given-names>S.</given-names></name> <name><surname>Tripathi</surname><given-names>V.</given-names></name> <name><surname>Bharadwaj</surname><given-names>A.</given-names></name></person-group> (<year>2025</year>). <article-title>Unravelling the secrets of soil microbiome and climate change for sustainable agroecosystems</article-title>. <source>Folia Microbiol.</source> <volume>70</volume>, <fpage>19</fpage>&#x2013;<lpage>40</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s12223-024-01194-9</pub-id></mixed-citation></ref>
<ref id="ref60"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Kaya</surname><given-names>C.</given-names></name></person-group> (<year>2024</year>). <article-title>Microbial modulation of hormone signaling, proteomic dynamics, and metabolomics in plant drought adaptation</article-title>. <source>Food Energy Secur.</source> <volume>13</volume>:<fpage>e513</fpage>. doi: <pub-id pub-id-type="doi">10.1002/fes3.513</pub-id></mixed-citation></ref>
<ref id="ref61"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Khan</surname><given-names>N.</given-names></name> <name><surname>Ali</surname><given-names>S.</given-names></name> <name><surname>Shahid</surname><given-names>M. A.</given-names></name> <name><surname>Mustafa</surname><given-names>A.</given-names></name> <name><surname>Sayyed</surname><given-names>R. Z.</given-names></name> <name><surname>Cur&#x00E1;</surname><given-names>J. A.</given-names></name></person-group> (<year>2021</year>). <article-title>Insights into the interactions among roots, rhizosphere, and rhizobacteria for improving plant growth and tolerance to abiotic stresses: a review</article-title>. <source>Cells</source> <volume>10</volume>:<fpage>1551</fpage>. doi: <pub-id pub-id-type="doi">10.3390/cells10061551</pub-id>, <pub-id pub-id-type="pmid">34205352</pub-id></mixed-citation></ref>
<ref id="ref62"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Khan</surname><given-names>N.</given-names></name> <name><surname>Bano</surname><given-names>A.</given-names></name> <name><surname>Ali</surname><given-names>S.</given-names></name> <name><surname>Babar</surname><given-names>M. A.</given-names></name></person-group> (<year>2020</year>). <article-title>Crosstalk amongst phytohormones from planta and PGPR under biotic and abiotic stresses</article-title>. <source>Plant Growth Regul.</source> <volume>90</volume>, <fpage>189</fpage>&#x2013;<lpage>203</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s10725-020-00571-x</pub-id></mixed-citation></ref>
<ref id="ref63"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Khan</surname><given-names>F.</given-names></name> <name><surname>Siddique</surname><given-names>A. B.</given-names></name> <name><surname>Shabala</surname><given-names>S.</given-names></name> <name><surname>Zhou</surname><given-names>M.</given-names></name> <name><surname>Zhao</surname><given-names>C.</given-names></name></person-group> (<year>2023</year>). <article-title>Phosphorus plays key roles in regulating plants&#x2019; physiological responses to abiotic stresses</article-title>. <source>Plants</source> <volume>12</volume>:<fpage>2861</fpage>. doi: <pub-id pub-id-type="doi">10.3390/plants12152861</pub-id>, <pub-id pub-id-type="pmid">37571014</pub-id></mixed-citation></ref>
<ref id="ref64"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Kong</surname><given-names>W.</given-names></name> <name><surname>Meldgin</surname><given-names>D. R.</given-names></name> <name><surname>Collins</surname><given-names>J. J.</given-names></name> <name><surname>Lu</surname><given-names>T.</given-names></name></person-group> (<year>2018</year>). <article-title>Designing microbial consortia with defined social interactions</article-title>. <source>Nat. Chem. Biol.</source> <volume>14</volume>, <fpage>821</fpage>&#x2013;<lpage>829</lpage>. doi: <pub-id pub-id-type="doi">10.1038/s41589-018-0091-7</pub-id>, <pub-id pub-id-type="pmid">29942078</pub-id></mixed-citation></ref>
<ref id="ref65"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Kotze</surname><given-names>C.</given-names></name> <name><surname>Van Niekerk</surname><given-names>J.</given-names></name> <name><surname>Mostert</surname><given-names>L.</given-names></name> <name><surname>Halleen</surname><given-names>F.</given-names></name> <name><surname>Fourie</surname><given-names>P.</given-names></name></person-group> (<year>2011</year>). <article-title>Evaluation of biocontrol agents for grapevine pruning wound protection against trunk pathogen infection</article-title>. <source>Phytopathol. Mediterr.</source> <volume>50</volume>, <fpage>S247</fpage>&#x2013;<lpage>S263</lpage>. doi: <pub-id pub-id-type="doi">10.14601/Phytopathol_Mediterr-8960</pub-id></mixed-citation></ref>
<ref id="ref66"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Kour</surname><given-names>D.</given-names></name> <name><surname>Kour</surname><given-names>H.</given-names></name> <name><surname>Khan</surname><given-names>S. S.</given-names></name> <name><surname>Khan</surname><given-names>R. T.</given-names></name> <name><surname>Bhardwaj</surname><given-names>M.</given-names></name> <name><surname>Kailoo</surname><given-names>S.</given-names></name> <etal/></person-group>. (<year>2023</year>). <article-title>Biodiversity and functional attributes of rhizospheric microbiomes: potential tools for sustainable agriculture</article-title>. <source>Curr. Microbiol.</source> <volume>80</volume>:<fpage>192</fpage>. doi: <pub-id pub-id-type="doi">10.1007/s00284-023-03300-5</pub-id>, <pub-id pub-id-type="pmid">37101055</pub-id></mixed-citation></ref>
<ref id="ref67"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Kumar</surname><given-names>H.</given-names></name> <name><surname>Verma</surname><given-names>S.</given-names></name> <name><surname>Rupali</surname><given-names>J. S.</given-names></name> <name><surname>Sagar</surname><given-names>D.</given-names></name></person-group> (<year>2024</year>). <article-title>A comprehensive review on entomovectoring in agroecosystem</article-title>. <source>J. Sci. Res. Rep.</source> <volume>30</volume>, <fpage>190</fpage>&#x2013;<lpage>200</lpage>. doi: <pub-id pub-id-type="doi">10.9734/jsrr/2024/v30i82239</pub-id></mixed-citation></ref>
<ref id="ref68"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Kumawat</surname><given-names>K. C.</given-names></name> <name><surname>Sharma</surname><given-names>B.</given-names></name> <name><surname>Nagpal</surname><given-names>S.</given-names></name> <name><surname>Kumar</surname><given-names>A.</given-names></name> <name><surname>Tiwari</surname><given-names>S.</given-names></name> <name><surname>Nair</surname><given-names>R. M.</given-names></name></person-group> (<year>2023</year>). <article-title>Plant growth-promoting rhizobacteria: salt stress alleviators to improve crop productivity for sustainable agriculture development</article-title>. <source>Front. Plant Sci.</source> <volume>13</volume>:<fpage>1101862</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fpls.2022.1101862</pub-id>, <pub-id pub-id-type="pmid">36714780</pub-id></mixed-citation></ref>
<ref id="ref69"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Lahlali</surname><given-names>R.</given-names></name> <name><surname>Ezrari</surname><given-names>S.</given-names></name> <name><surname>Radouane</surname><given-names>N.</given-names></name> <name><surname>Kenfaoui</surname><given-names>J.</given-names></name> <name><surname>Esmaeel</surname><given-names>Q.</given-names></name> <name><surname>El Hamss</surname><given-names>H.</given-names></name> <etal/></person-group>. (<year>2022</year>). <article-title>Biological control of plant pathogens: a global perspective</article-title>. <source>Microorganisms</source> <volume>10</volume>:<fpage>596</fpage>. doi: <pub-id pub-id-type="doi">10.3390/microorganisms10030596</pub-id>, <pub-id pub-id-type="pmid">35336171</pub-id></mixed-citation></ref>
<ref id="ref70"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Leal</surname><given-names>C.</given-names></name> <name><surname>Eichmeier</surname><given-names>A.</given-names></name> <name><surname>&#x0160;t&#x016F;skov&#x00E1;</surname><given-names>K.</given-names></name> <name><surname>Armengol</surname><given-names>J.</given-names></name> <name><surname>Bujanda</surname><given-names>R.</given-names></name> <name><surname>Fontaine</surname><given-names>F.</given-names></name> <etal/></person-group>. (<year>2024</year>). <article-title>Establishment of biocontrol agents and their impact on rhizosphere microbiome and induced grapevine defenses are highly soil-dependent</article-title>. <source>Phytobiomes J.</source> <volume>8</volume>, <fpage>111</fpage>&#x2013;<lpage>127</lpage>. doi: <pub-id pub-id-type="doi">10.1094/PBIOMES-08-23-0077-R</pub-id></mixed-citation></ref>
<ref id="ref71"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Lee D&#x00ED;az</surname><given-names>A. S.</given-names></name> <name><surname>Macheda</surname><given-names>D.</given-names></name> <name><surname>Saha</surname><given-names>H.</given-names></name> <name><surname>Ploll</surname><given-names>U.</given-names></name> <name><surname>Orine</surname><given-names>D.</given-names></name> <name><surname>Biere</surname><given-names>A.</given-names></name></person-group> (<year>2021</year>). <article-title>Tackling the context-dependency of microbial-induced resistance</article-title>. <source>Agronomy</source> <volume>11</volume>:<fpage>1293</fpage>. doi: <pub-id pub-id-type="doi">10.3390/agronomy11071293</pub-id></mixed-citation></ref>
<ref id="ref72"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Li</surname><given-names>J.</given-names></name> <name><surname>Wang</surname><given-names>C.</given-names></name> <name><surname>Liang</surname><given-names>W.</given-names></name> <name><surname>Liu</surname><given-names>S.</given-names></name></person-group> (<year>2021</year>). <article-title>Rhizosphere microbiome: the emerging barrier in plant-pathogen interactions</article-title>. <source>Front. Microbiol.</source> <volume>12</volume>:<fpage>772420</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fmicb.2021.772420</pub-id>, <pub-id pub-id-type="pmid">34777326</pub-id></mixed-citation></ref>
<ref id="ref73"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Liao</surname><given-names>H.</given-names></name> <name><surname>Hao</surname><given-names>X.</given-names></name> <name><surname>Li</surname><given-names>Y.</given-names></name> <name><surname>Ma</surname><given-names>S.</given-names></name> <name><surname>Gao</surname><given-names>S.</given-names></name> <name><surname>Cai</surname><given-names>P.</given-names></name> <etal/></person-group>. (<year>2024</year>). <article-title>Protists regulate microbially mediated organic carbon turnover in soil aggregates</article-title>. <source>Glob. Change Biol.</source> <volume>30</volume>:<fpage>e17102</fpage>. doi: <pub-id pub-id-type="doi">10.1111/gcb.17102</pub-id>, <pub-id pub-id-type="pmid">38273557</pub-id></mixed-citation></ref>
<ref id="ref74"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Lucas</surname><given-names>J. A.</given-names></name> <name><surname>Hawkins</surname><given-names>N. J.</given-names></name> <name><surname>Fraaije</surname><given-names>B. A.</given-names></name></person-group> (<year>2015</year>). <article-title>The evolution of fungicide resistance</article-title>. <source>Adv. Appl. Microbiol.</source> <volume>90</volume>, <fpage>29</fpage>&#x2013;<lpage>92</lpage>. doi: <pub-id pub-id-type="doi">10.1016/bs.aambs.2014.09.001</pub-id>, <pub-id pub-id-type="pmid">25596029</pub-id></mixed-citation></ref>
<ref id="ref75"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Mamun</surname><given-names>A. A.</given-names></name> <name><surname>Neumann</surname><given-names>G.</given-names></name> <name><surname>Moradtalab</surname><given-names>N.</given-names></name> <name><surname>Ahmed</surname><given-names>A.</given-names></name> <name><surname>Dupuis</surname><given-names>B.</given-names></name> <name><surname>Darbon</surname><given-names>G.</given-names></name> <etal/></person-group>. (<year>2024</year>). <article-title>Microbial consortia versus single-strain inoculants as drought stress protectants in potato affected by the form of N supply</article-title>. <source>Horticulturae</source> <volume>10</volume>:<fpage>102</fpage>. doi: <pub-id pub-id-type="doi">10.3390/horticulturae10010102</pub-id></mixed-citation></ref>
<ref id="ref76"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Mannaa</surname><given-names>M.</given-names></name> <name><surname>Han</surname><given-names>G.</given-names></name> <name><surname>Jung</surname><given-names>H.</given-names></name> <name><surname>Park</surname><given-names>J.</given-names></name> <name><surname>Kim</surname><given-names>J. C.</given-names></name> <name><surname>Park</surname><given-names>A. R.</given-names></name> <etal/></person-group>. (<year>2023</year>). <article-title><italic>Aureobasidium pullulans</italic> treatment mitigates drought stress in <italic>Abies koreana</italic> via rhizosphere microbiome modulation</article-title>. <source>Plants</source> <volume>12</volume>:<fpage>3653</fpage>. doi: <pub-id pub-id-type="doi">10.3390/plants12203653</pub-id>, <pub-id pub-id-type="pmid">37896116</pub-id></mixed-citation></ref>
<ref id="ref77"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Marzouk</surname><given-names>S. H.</given-names></name> <name><surname>Kwaslema</surname><given-names>D. R.</given-names></name> <name><surname>Omar</surname><given-names>M. M.</given-names></name> <name><surname>Mohamed</surname><given-names>S. H.</given-names></name></person-group> (<year>2025</year>). <article-title>Harnessing the power of soil microbes: their dual impact in integrated nutrient management and mediating climate stress for sustainable rice crop production&#x2014;a systematic review</article-title>. <source>Heliyon</source> <volume>11</volume>:<fpage>e41158</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.heliyon.2024.e41158</pub-id>, <pub-id pub-id-type="pmid">39758363</pub-id></mixed-citation></ref>
<ref id="ref78"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Massart</surname><given-names>S.</given-names></name> <name><surname>Martinez-Medina</surname><given-names>M.</given-names></name> <name><surname>Jijakli</surname><given-names>M. H.</given-names></name></person-group> (<year>2015</year>). <article-title>Biological control in the microbiome era: challenges and opportunities</article-title>. <source>Biol. Control</source> <volume>89</volume>, <fpage>98</fpage>&#x2013;<lpage>108</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.biocontrol.2015.06.003</pub-id></mixed-citation></ref>
<ref id="ref79"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Mazzola</surname><given-names>M.</given-names></name></person-group> (<year>2002</year>). <article-title>Mechanisms of natural soil suppressiveness to soilborne diseases</article-title>. <source>Antonie Van Leeuwenhoek</source> <volume>81</volume>, <fpage>557</fpage>&#x2013;<lpage>564</lpage>. doi: <pub-id pub-id-type="doi">10.1023/A:1020557523557</pub-id></mixed-citation></ref>
<ref id="ref80"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Mehmood</surname><given-names>N.</given-names></name> <name><surname>Saeed</surname><given-names>M.</given-names></name> <name><surname>Zafarullah</surname><given-names>S.</given-names></name> <name><surname>Hyder</surname><given-names>S.</given-names></name> <name><surname>Rizvi</surname><given-names>Z. F.</given-names></name> <name><surname>Gondal</surname><given-names>A. S.</given-names></name> <etal/></person-group>. (<year>2023</year>). <article-title>Multifaceted impacts of plant-beneficial <italic>Pseudomonas</italic> spp. in managing various plant diseases and crop yield improvement</article-title>. <source>ACS Omega</source> <volume>8</volume>, <fpage>22296</fpage>&#x2013;<lpage>22315</lpage>. doi: <pub-id pub-id-type="doi">10.1021/acsomega.3c00870</pub-id>, <pub-id pub-id-type="pmid">37396244</pub-id></mixed-citation></ref>
<ref id="ref81"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Men&#x00E9;ndez-Ca&#x00F1;amares</surname><given-names>S.</given-names></name> <name><surname>Bl&#x00E1;zquez</surname><given-names>A.</given-names></name> <name><surname>Albertos</surname><given-names>I.</given-names></name> <name><surname>Poveda</surname><given-names>J.</given-names></name> <name><surname>D&#x00ED;ez-M&#x00E9;ndez</surname><given-names>A.</given-names></name></person-group> (<year>2024</year>). <article-title>Probiotic <italic>Bacillus subtilis</italic> SB8 and edible coatings for sustainable fungal disease management in strawberry</article-title>. <source>Biol. Control</source> <volume>196</volume>:<fpage>105572</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.biocontrol.2024.105572</pub-id></mixed-citation></ref>
<ref id="ref82"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Mesguida</surname><given-names>O.</given-names></name> <name><surname>Haidar</surname><given-names>R.</given-names></name> <name><surname>Yacoub</surname><given-names>A.</given-names></name> <name><surname>Dreux-Zigha</surname><given-names>A.</given-names></name> <name><surname>Berthon</surname><given-names>J. Y.</given-names></name> <name><surname>Guyoneaud</surname><given-names>R.</given-names></name> <etal/></person-group>. (<year>2023</year>). <article-title>Microbial biological control of fungi associated with grapevine trunk diseases: a review of strain diversity, modes of action, and advantages and limits of current strategies</article-title>. <source>J. Fungi</source> <volume>9</volume>:<fpage>638</fpage>. doi: <pub-id pub-id-type="doi">10.3390/jof9060638</pub-id>, <pub-id pub-id-type="pmid">37367574</pub-id></mixed-citation></ref>
<ref id="ref83"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Minchev</surname><given-names>Z.</given-names></name> <name><surname>Kostenko</surname><given-names>O.</given-names></name> <name><surname>Soler</surname><given-names>R.</given-names></name> <name><surname>Pozo</surname><given-names>M. J.</given-names></name></person-group> (<year>2021</year>). <article-title>Microbial consortia for effective biocontrol of root and foliar diseases in tomato</article-title>. <source>Front. Plant Sci.</source> <volume>12</volume>:<fpage>756368</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fpls.2021.756368</pub-id>, <pub-id pub-id-type="pmid">34804094</pub-id></mixed-citation></ref>
<ref id="ref84"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Mon</surname><given-names>Y. Y.</given-names></name> <name><surname>Bidabadi</surname><given-names>S. S.</given-names></name> <name><surname>Oo</surname><given-names>K. S.</given-names></name> <name><surname>Zheng</surname><given-names>S. J.</given-names></name></person-group> (<year>2021</year>). <article-title>The antagonistic mechanism of rhizosphere microbes and endophytes on the interaction between banana and <italic>fusarium oxysporum</italic> f. sp. <italic>cubense</italic></article-title>. <source>Physiol. Mol. Plant Pathol.</source> <volume>116</volume>:<fpage>101733</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.pmpp.2021.101733</pub-id></mixed-citation></ref>
<ref id="ref85"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Mona</surname><given-names>S. A.</given-names></name> <name><surname>Hashem</surname><given-names>A.</given-names></name> <name><surname>Abd_Allah</surname><given-names>E. F.</given-names></name> <name><surname>Alqarawi</surname><given-names>A. A.</given-names></name> <name><surname>Soliman</surname><given-names>D. W. K.</given-names></name> <name><surname>Wirth</surname><given-names>S.</given-names></name> <etal/></person-group>. (<year>2017</year>). <article-title>Increased resistance of drought by <italic>Trichoderma harzianum</italic> fungal treatment correlates with increased secondary metabolites and proline content</article-title>. <source>J. Integr. Agric.</source> <volume>16</volume>, <fpage>1751</fpage>&#x2013;<lpage>1757</lpage>. doi: <pub-id pub-id-type="doi">10.1016/S2095-3119(17)61695-2</pub-id></mixed-citation></ref>
<ref id="ref86"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Moussa</surname><given-names>S.</given-names></name> <name><surname>Iasur Kruh</surname><given-names>L.</given-names></name></person-group> (<year>2025</year>). <article-title>Balancing nature and nurture: the role of biocontrol agents in shaping plant microbiomes for sustainable agriculture</article-title>. <source>Microorganisms</source> <volume>13</volume>:<fpage>323</fpage>. doi: <pub-id pub-id-type="doi">10.3390/microorganisms13020323</pub-id>, <pub-id pub-id-type="pmid">40005690</pub-id></mixed-citation></ref>
<ref id="ref87"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Muhammad</surname><given-names>A.</given-names></name> <name><surname>Kong</surname><given-names>X.</given-names></name> <name><surname>Zheng</surname><given-names>S.</given-names></name> <name><surname>Bai</surname><given-names>N.</given-names></name> <name><surname>Li</surname><given-names>L.</given-names></name> <name><surname>Khan</surname><given-names>M. H. U.</given-names></name> <etal/></person-group>. (<year>2024</year>). <article-title>Exploring plant&#x2013;microbe interactions in adapting to abiotic stress under climate change: a review</article-title>. <source>Front. Plant Sci.</source> <volume>15</volume>:<fpage>1482739</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fpls.2024.1482739</pub-id>, <pub-id pub-id-type="pmid">39619840</pub-id></mixed-citation></ref>
<ref id="ref88"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Munir</surname><given-names>N.</given-names></name> <name><surname>Hanif</surname><given-names>M.</given-names></name> <name><surname>Abideen</surname><given-names>Z.</given-names></name> <name><surname>Sohail</surname><given-names>M.</given-names></name> <name><surname>El-Keblawy</surname><given-names>A.</given-names></name> <name><surname>Radicetti</surname><given-names>E.</given-names></name> <etal/></person-group>. (<year>2022</year>). <article-title>Mechanisms and strategies of plant microbiome interactions to mitigate abiotic stresses</article-title>. <source>Agronomy</source> <volume>12</volume>:<fpage>2069</fpage>. doi: <pub-id pub-id-type="doi">10.3390/agronomy12092069</pub-id></mixed-citation></ref>
<ref id="ref89"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Negre Rodr&#x00ED;guez</surname><given-names>M.</given-names></name> <name><surname>Pioppi</surname><given-names>A.</given-names></name> <name><surname>Kov&#x00E1;cs</surname><given-names>&#x00C1;. T.</given-names></name></person-group> (<year>2025</year>). <article-title>The role of plant host genetics in shaping the composition and functionality of rhizosphere microbiomes</article-title>. <source>mSystems</source> <volume>10</volume>:<fpage>e00041-24</fpage>. doi: <pub-id pub-id-type="doi">10.1128/msystems.00041-24</pub-id>, <pub-id pub-id-type="pmid">40643236</pub-id></mixed-citation></ref>
<ref id="ref90"><mixed-citation publication-type="book"><person-group person-group-type="author"><name><surname>Ni</surname><given-names>L.</given-names></name> <name><surname>Punja</surname><given-names>Z. K.</given-names></name></person-group> (<year>2020</year>). &#x201C;<chapter-title>Management of fungal diseases on cucumber (<italic>Cucumis sativus</italic> L.) and tomato (<italic>Solanum lycopersicum</italic> L.) crops in greenhouses using <italic>Bacillus subtilis</italic></chapter-title>&#x201D; in <source>Bacilli and Agrobiotechnology: Phytostimulation and Biocontrol</source>. eds. <person-group person-group-type="editor"><name><surname>Islam</surname><given-names>M.</given-names></name> <name><surname>Rahman</surname><given-names>M.</given-names></name> <name><surname>Pandey</surname><given-names>P.</given-names></name> <name><surname>Boehme</surname><given-names>M.</given-names></name> <name><surname>Haesaert</surname><given-names>G.</given-names></name></person-group> (<publisher-loc>Cham</publisher-loc>: <publisher-name>Springer</publisher-name>), <fpage>1</fpage>&#x2013;<lpage>28</lpage>.</mixed-citation></ref>
<ref id="ref91"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Nie</surname><given-names>W.</given-names></name> <name><surname>He</surname><given-names>Q.</given-names></name> <name><surname>Guo</surname><given-names>H.</given-names></name> <name><surname>Zhang</surname><given-names>W.</given-names></name> <name><surname>Ma</surname><given-names>L.</given-names></name> <name><surname>Li</surname><given-names>J.</given-names></name> <etal/></person-group>. (<year>2024</year>). <article-title>Arbuscular mycorrhizal fungi: boosting crop resilience to environmental stresses</article-title>. <source>Microorganisms</source> <volume>12</volume>:<fpage>2448</fpage>. doi: <pub-id pub-id-type="doi">10.3390/microorganisms12122448</pub-id>, <pub-id pub-id-type="pmid">39770651</pub-id></mixed-citation></ref>
<ref id="ref92"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Nivetha</surname><given-names>N.</given-names></name> <name><surname>Asha</surname><given-names>A. D.</given-names></name> <name><surname>Krishna</surname><given-names>G. K.</given-names></name> <name><surname>Chinnusamy</surname><given-names>V.</given-names></name> <name><surname>Paul</surname><given-names>S.</given-names></name></person-group> (<year>2024</year>). <article-title>Rhizobacteria <italic>Bacillus</italic> spp. mitigate osmotic stress and improve seed germination in mustard by regulating osmolyte and plant hormone signaling</article-title>. <source>Physiol. Plant.</source> <volume>176</volume>:<fpage>e14202</fpage>. doi: <pub-id pub-id-type="doi">10.1111/ppl.14202</pub-id>, <pub-id pub-id-type="pmid">38356406</pub-id></mixed-citation></ref>
<ref id="ref93"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Obiazikwor</surname><given-names>O. H.</given-names></name> <name><surname>Shah</surname><given-names>A.</given-names></name> <name><surname>Hardy</surname><given-names>G. S. J.</given-names></name> <name><surname>Bayliss</surname><given-names>K.</given-names></name></person-group> (<year>2025</year>). <article-title>The rhizosphere microbiome can sustainably protect field-grown tomato crops against soil-borne pathogens and plant parasitic nematodes</article-title>. <source>Can. J. Plant Pathol.</source> <volume>47</volume>, <fpage>423</fpage>&#x2013;<lpage>436</lpage>. doi: <pub-id pub-id-type="doi">10.1080/07060661.2025.2477644</pub-id></mixed-citation></ref>
<ref id="ref94"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Offre</surname><given-names>P.</given-names></name> <name><surname>Spang</surname><given-names>A.</given-names></name> <name><surname>Schleper</surname><given-names>C.</given-names></name></person-group> (<year>2013</year>). <article-title>Archaea in biogeochemical cycles</article-title>. <source>Ann. Rev. Microbiol.</source> <volume>67</volume>, <fpage>437</fpage>&#x2013;<lpage>457</lpage>. doi: <pub-id pub-id-type="doi">10.1146/annurev-micro-092412-155614</pub-id>, <pub-id pub-id-type="pmid">23808334</pub-id></mixed-citation></ref>
<ref id="ref95"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Orozco-Mosqueda</surname><given-names>M. D. C.</given-names></name> <name><surname>Santoyo</surname><given-names>G.</given-names></name> <name><surname>Glick</surname><given-names>B. R.</given-names></name></person-group> (<year>2023</year>). <article-title>Recent advances in the bacterial phytohormone modulation of plant growth</article-title>. <source>Plants</source> <volume>12</volume>:<fpage>606</fpage>. doi: <pub-id pub-id-type="doi">10.3390/plants12030606</pub-id>, <pub-id pub-id-type="pmid">36771689</pub-id></mixed-citation></ref>
<ref id="ref96"><mixed-citation publication-type="book"><person-group person-group-type="author"><name><surname>Panpatte</surname><given-names>D. G.</given-names></name> <name><surname>Jhala</surname><given-names>Y. K.</given-names></name> <name><surname>Shelat</surname><given-names>H. N.</given-names></name> <name><surname>Vyas</surname><given-names>R. V.</given-names></name></person-group> (<year>2016</year>). &#x201C;<chapter-title><italic>Pseudomonas fluorescens</italic>: a promising biocontrol agent and PGPR for sustainable agriculture</chapter-title>&#x201D; in <source>Microbial Inoculants in Sustainable Agricultural Productivity</source>. eds. <person-group person-group-type="editor"><name><surname>Singh</surname><given-names>D. P.</given-names></name> <name><surname>Singh</surname><given-names>H. B.</given-names></name> <name><surname>Prabha</surname><given-names>R.</given-names></name></person-group> (<publisher-loc>New Delhi</publisher-loc>: <publisher-name>Springer</publisher-name>), <fpage>257</fpage>&#x2013;<lpage>270</lpage>.</mixed-citation></ref>
<ref id="ref97"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Park</surname><given-names>I.</given-names></name> <name><surname>Seo</surname><given-names>Y. S.</given-names></name> <name><surname>Mannaa</surname><given-names>M.</given-names></name></person-group> (<year>2023</year>). <article-title>Recruitment of the rhizo-microbiome army: assembly determinants and engineering of the rhizosphere microbiome as a key to unlocking plant potential</article-title>. <source>Front. Microbiol.</source> <volume>14</volume>:<fpage>1163832</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fmicb.2023.1163832</pub-id>, <pub-id pub-id-type="pmid">37213524</pub-id></mixed-citation></ref>
<ref id="ref98"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Pathak</surname><given-names>V. M.</given-names></name> <name><surname>Verma</surname><given-names>V. K.</given-names></name> <name><surname>Rawat</surname><given-names>B. S.</given-names></name> <name><surname>Kaur</surname><given-names>B.</given-names></name> <name><surname>Babu</surname><given-names>N.</given-names></name> <name><surname>Sharma</surname><given-names>A.</given-names></name> <etal/></person-group>. (<year>2022</year>). <article-title>Current status of pesticide effects on environment, human health and its eco-friendly management as bioremediation: a comprehensive review</article-title>. <source>Front. Microbiol.</source> <volume>13</volume>:<fpage>962619</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fmicb.2022.962619</pub-id></mixed-citation></ref>
<ref id="ref99"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Pearson</surname><given-names>D. E.</given-names></name> <name><surname>Callaway</surname><given-names>R. M.</given-names></name></person-group> (<year>2003</year>). <article-title>Indirect effects of host-specific biological control agents</article-title>. <source>Trends Ecol. Evol.</source> <volume>18</volume>, <fpage>456</fpage>&#x2013;<lpage>461</lpage>. doi: <pub-id pub-id-type="doi">10.1016/S0169-5347(03)00188-5</pub-id></mixed-citation></ref>
<ref id="ref100"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Pearson</surname><given-names>D. E.</given-names></name> <name><surname>Callaway</surname><given-names>R. M.</given-names></name></person-group> (<year>2005</year>). <article-title>Indirect nontarget effects of host-specific biological control agents: implications for biological control</article-title>. <source>Biol. Control</source> <volume>35</volume>, <fpage>288</fpage>&#x2013;<lpage>298</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.biocontrol.2005.05.011</pub-id></mixed-citation></ref>
<ref id="ref101"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Peng</surname><given-names>J.</given-names></name> <name><surname>Mavrodi</surname><given-names>D. V.</given-names></name> <name><surname>Li</surname><given-names>J.</given-names></name> <name><surname>Egan</surname><given-names>S.</given-names></name> <name><surname>Zhang</surname><given-names>H.</given-names></name> <name><surname>Fan</surname><given-names>X.</given-names></name> <etal/></person-group>. (<year>2025</year>). <article-title><italic>Stenotrophomonas maltophilia</italic> impedes <italic>Bacillus</italic> biocontrol of tomato wilt disease by degrading its lipopeptide antibiotics</article-title>. <source>ISME J.</source> <volume>19</volume>:<fpage>wraf210</fpage>. doi: <pub-id pub-id-type="doi">10.1093/ismejo/wraf210</pub-id></mixed-citation></ref>
<ref id="ref102"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Philippot</surname><given-names>L.</given-names></name> <name><surname>Raaijmakers</surname><given-names>J. M.</given-names></name> <name><surname>Lemanceau</surname><given-names>P.</given-names></name> <name><surname>Van Der Putten</surname><given-names>W. H.</given-names></name></person-group> (<year>2013</year>). <article-title>Going back to the roots: the microbial ecology of the rhizosphere</article-title>. <source>Nat. Rev. Microbiol.</source> <volume>11</volume>, <fpage>789</fpage>&#x2013;<lpage>799</lpage>. doi: <pub-id pub-id-type="doi">10.1038/nrmicro3109</pub-id>, <pub-id pub-id-type="pmid">24056930</pub-id></mixed-citation></ref>
<ref id="ref103"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Pieterse</surname><given-names>C. M.</given-names></name> <name><surname>Zamioudis</surname><given-names>C.</given-names></name> <name><surname>Berendsen</surname><given-names>R. L.</given-names></name> <name><surname>Weller</surname><given-names>D. M.</given-names></name> <name><surname>Van Wees</surname><given-names>S. C.</given-names></name> <name><surname>Bakker</surname><given-names>P. A.</given-names></name></person-group> (<year>2014</year>). <article-title>Induced systemic resistance by beneficial microbes</article-title>. <source>Annu. Rev. Phytopathol.</source> <volume>52</volume>, <fpage>347</fpage>&#x2013;<lpage>375</lpage>. doi: <pub-id pub-id-type="doi">10.1146/annurev-phyto-082712-102340</pub-id>, <pub-id pub-id-type="pmid">24906124</pub-id></mixed-citation></ref>
<ref id="ref104"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Ploetz</surname><given-names>R. C.</given-names></name></person-group> (<year>2015</year>). <article-title>Management of <italic>Fusarium</italic> wilt of banana: a review with special reference to tropical race 4</article-title>. <source>Crop Prot.</source> <volume>73</volume>, <fpage>7</fpage>&#x2013;<lpage>15</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.cropro.2015.01.007</pub-id></mixed-citation></ref>
<ref id="ref105"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Poveda</surname><given-names>J.</given-names></name></person-group> (<year>2021</year>). <article-title>Insect frass in the development of sustainable agriculture: a review</article-title>. <source>Agron. Sustain. Dev.</source> <volume>41</volume>:<fpage>5</fpage>. doi: <pub-id pub-id-type="doi">10.1007/s13593-020-00656-x</pub-id></mixed-citation></ref>
<ref id="ref106"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Preece</surname><given-names>C.</given-names></name> <name><surname>Pe&#x00F1;uelas</surname><given-names>J.</given-names></name></person-group> (<year>2016</year>). <article-title>Rhizodeposition under drought and consequences for soil communities and ecosystem resilience</article-title>. <source>Plant Soil</source> <volume>409</volume>, <fpage>1</fpage>&#x2013;<lpage>17</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s11104-016-3090-z</pub-id></mixed-citation></ref>
<ref id="ref107"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Prosser</surname><given-names>J. I.</given-names></name></person-group> (<year>2012</year>). <article-title>Ecosystem processes and interactions in a morass of diversity</article-title>. <source>FEMS Microbiol. Ecol.</source> <volume>81</volume>, <fpage>507</fpage>&#x2013;<lpage>519</lpage>. doi: <pub-id pub-id-type="doi">10.1111/j.1574-6941.2012.01435.x</pub-id>, <pub-id pub-id-type="pmid">22715974</pub-id></mixed-citation></ref>
<ref id="ref108"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Raaijmakers</surname><given-names>J. M.</given-names></name> <name><surname>Mazzola</surname><given-names>M.</given-names></name></person-group> (<year>2012</year>). <article-title>Diversity and natural functions of antibiotics produced by beneficial and plant pathogenic bacteria</article-title>. <source>Annu. Rev. Phytopathol.</source> <volume>50</volume>, <fpage>403</fpage>&#x2013;<lpage>424</lpage>. doi: <pub-id pub-id-type="doi">10.1146/annurev-phyto-081211-172908</pub-id>, <pub-id pub-id-type="pmid">22681451</pub-id></mixed-citation></ref>
<ref id="ref109"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Rabari</surname><given-names>A.</given-names></name> <name><surname>Ruparelia</surname><given-names>J.</given-names></name> <name><surname>Jha</surname><given-names>C. K.</given-names></name> <name><surname>Sayyed</surname><given-names>R. Z.</given-names></name> <name><surname>Mitra</surname><given-names>D.</given-names></name> <name><surname>Priyadarshini</surname><given-names>A.</given-names></name> <etal/></person-group>. (<year>2023</year>). <article-title>Articulating beneficial rhizobacteria-mediated plant defenses through induced systemic resistance: a review</article-title>. <source>Pedosphere</source> <volume>33</volume>, <fpage>556</fpage>&#x2013;<lpage>566</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.pedsph.2022.10.003</pub-id></mixed-citation></ref>
<ref id="ref110"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Ramegowda</surname><given-names>V.</given-names></name> <name><surname>Senthil-Kumar</surname><given-names>M.</given-names></name></person-group> (<year>2015</year>). <article-title>The interactive effects of simultaneous biotic and abiotic stresses on plants: mechanistic understanding from drought and pathogen combination</article-title>. <source>J. Plant Physiol.</source> <volume>176</volume>, <fpage>47</fpage>&#x2013;<lpage>54</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.jplph.2014.11.008</pub-id>, <pub-id pub-id-type="pmid">25546584</pub-id></mixed-citation></ref>
<ref id="ref111"><mixed-citation publication-type="book"><person-group person-group-type="author"><name><surname>Rawal</surname><given-names>H.</given-names></name> <name><surname>Nautiyal</surname><given-names>Y.</given-names></name> <name><surname>Sharma</surname><given-names>B.</given-names></name> <name><surname>Tiwari</surname><given-names>S.</given-names></name></person-group> (<year>2024</year>). &#x201C;<chapter-title>Microbial inoculants and their role in abiotic stress management</chapter-title>&#x201D; in <source>Microbial Inoculants: Applications for Sustainable Agriculture</source>. eds. <person-group person-group-type="editor"><name><surname>Chaudhary</surname><given-names>P.</given-names></name> <name><surname>Chaudhary</surname><given-names>A.</given-names></name></person-group> (<publisher-loc>Singapore</publisher-loc>: <publisher-name>Springer Nature</publisher-name>), <fpage>163</fpage>&#x2013;<lpage>201</lpage>.</mixed-citation></ref>
<ref id="ref112"><mixed-citation publication-type="book"><person-group person-group-type="author"><name><surname>Rigobelo</surname><given-names>E. C.</given-names></name></person-group> (<year>2025</year>). &#x201C;<chapter-title>Endophytic filamentous fungi as sources of metabolites for agricultural applications</chapter-title>&#x201D; in <source>Fungal Metabolites for Agricultural Applications: Biostimulation and Crop Protection by Fungal Biotechnology</source>. eds. <person-group person-group-type="editor"><name><surname>Poveda</surname><given-names>J.</given-names></name> <name><surname>Santamaria</surname><given-names>O.</given-names></name> <name><surname>Martin-Garcia</surname><given-names>J.</given-names></name></person-group> (<publisher-loc>Cham</publisher-loc>: <publisher-name>Springer Nature</publisher-name>), <fpage>1</fpage>&#x2013;<lpage>20</lpage>.</mixed-citation></ref>
<ref id="ref113"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Rigobelo</surname><given-names>E. C.</given-names></name> <name><surname>de Andrade</surname><given-names>L. A.</given-names></name> <name><surname>Santos</surname><given-names>C. H. B.</given-names></name> <name><surname>Frezarin</surname><given-names>E. T.</given-names></name> <name><surname>Sales</surname><given-names>L. R.</given-names></name> <name><surname>de Carvalho</surname><given-names>L. A. L.</given-names></name> <etal/></person-group>. (<year>2024</year>). <article-title>Effects of <italic>Trichoderma harzianum</italic> and <italic>Bacillus subtilis</italic> on the root and soil microbiomes of the soybean plant INTACTA RR2 PRO&#x2122;</article-title>. <source>Front. Plant Sci.</source> <volume>15</volume>:<fpage>1403160</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fpls.2024.1403160</pub-id></mixed-citation></ref>
<ref id="ref114"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Ristaino</surname><given-names>J. B.</given-names></name> <name><surname>Anderson</surname><given-names>P. K.</given-names></name> <name><surname>Bebber</surname><given-names>D. P.</given-names></name> <name><surname>Brauman</surname><given-names>K. A.</given-names></name> <name><surname>Cunniffe</surname><given-names>N. J.</given-names></name> <name><surname>Fedoroff</surname><given-names>N. V.</given-names></name> <etal/></person-group>. (<year>2021</year>). <article-title>The persistent threat of emerging plant disease pandemics to global food security</article-title>. <source>Proc. Natl. Acad. Sci. USA</source> <volume>118</volume>:<fpage>e2022239118</fpage>. doi: <pub-id pub-id-type="doi">10.1073/pnas.2022239118</pub-id></mixed-citation></ref>
<ref id="ref115"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Rizvi</surname><given-names>A.</given-names></name> <name><surname>Khan</surname><given-names>M. S.</given-names></name></person-group> (<year>2018</year>). <article-title>Heavy metal-induced oxidative damage and root morphology alterations of maize (<italic>Zea mays</italic> L.) plants and stress mitigation by metal-tolerant nitrogen-fixing <italic>Azotobacter chroococcum</italic></article-title>. <source>Ecotoxicol. Environ. Saf.</source> <volume>157</volume>, <fpage>9</fpage>&#x2013;<lpage>20</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.ecoenv.2018.03.063</pub-id>, <pub-id pub-id-type="pmid">29605647</pub-id></mixed-citation></ref>
<ref id="ref116"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Rohr</surname><given-names>J. R.</given-names></name> <name><surname>Barrett</surname><given-names>C. B.</given-names></name> <name><surname>Civitello</surname><given-names>D. J.</given-names></name> <name><surname>Craft</surname><given-names>M. E.</given-names></name> <name><surname>Delius</surname><given-names>B.</given-names></name> <name><surname>DeLeo</surname><given-names>G. A.</given-names></name> <etal/></person-group>. (<year>2019</year>). <article-title>Emerging human infectious diseases and the links to global food production</article-title>. <source>Nat. Sustain.</source> <volume>2</volume>, <fpage>445</fpage>&#x2013;<lpage>456</lpage>. doi: <pub-id pub-id-type="doi">10.1038/s41893-019-0293-3</pub-id></mixed-citation></ref>
<ref id="ref117"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Rolfe</surname><given-names>S. A.</given-names></name> <name><surname>Griffiths</surname><given-names>J.</given-names></name> <name><surname>Ton</surname><given-names>J.</given-names></name></person-group> (<year>2019</year>). <article-title>Crying out for help with root exudates: adaptive mechanisms by which stressed plants assemble health-promoting soil microbiomes</article-title>. <source>Curr. Opin. Microbiol.</source> <volume>49</volume>, <fpage>73</fpage>&#x2013;<lpage>82</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.mib.2019.10.003</pub-id>, <pub-id pub-id-type="pmid">31731229</pub-id></mixed-citation></ref>
<ref id="ref118"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Rossi</surname><given-names>C. A.</given-names></name> <name><surname>Marchetta</surname><given-names>E. J.</given-names></name> <name><surname>Kim</surname><given-names>J. H.</given-names></name> <name><surname>Castroverde</surname><given-names>C. D. M.</given-names></name></person-group> (<year>2023</year>). <article-title>Molecular regulation of the salicylic acid hormone pathway in plants under changing environmental conditions</article-title>. <source>Trends Biochem. Sci.</source> <volume>48</volume>, <fpage>699</fpage>&#x2013;<lpage>712</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.tibs.2023.05.004</pub-id>, <pub-id pub-id-type="pmid">37258325</pub-id></mixed-citation></ref>
<ref id="ref119"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Sackey</surname><given-names>O. K.</given-names></name> <name><surname>Feng</surname><given-names>N.</given-names></name> <name><surname>Mohammed</surname><given-names>Y. Z.</given-names></name> <name><surname>Dzou</surname><given-names>C. F.</given-names></name> <name><surname>Zheng</surname><given-names>D.</given-names></name> <name><surname>Zhao</surname><given-names>L.</given-names></name> <etal/></person-group>. (<year>2025</year>). <article-title>A comprehensive review on rice responses and tolerance to salt stress</article-title>. <source>Front. Plant Sci.</source> <volume>16</volume>:<fpage>1561280</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fpls.2025.1561280</pub-id>, <pub-id pub-id-type="pmid">40230606</pub-id></mixed-citation></ref>
<ref id="ref120"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Saleem</surname><given-names>M. A.</given-names></name> <name><surname>Khan</surname><given-names>A.</given-names></name> <name><surname>Tu</surname><given-names>J.</given-names></name> <name><surname>Huang</surname><given-names>W.</given-names></name> <name><surname>Liu</surname><given-names>Y.</given-names></name> <name><surname>Feng</surname><given-names>N.</given-names></name> <etal/></person-group>. (<year>2025</year>). <article-title>Salinity stress in rice: multilayered approaches for sustainable tolerance</article-title>. <source>Int. J. Mol. Sci.</source> <volume>26</volume>:<fpage>6025</fpage>. doi: <pub-id pub-id-type="doi">10.3390/ijms26136025</pub-id>, <pub-id pub-id-type="pmid">40649804</pub-id></mixed-citation></ref>
<ref id="ref121"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>S&#x00E1;nchez-Bayo</surname><given-names>F.</given-names></name> <name><surname>Wyckhuys</surname><given-names>K. A.</given-names></name></person-group> (<year>2019</year>). <article-title>Worldwide decline of the entomofauna: a review of its drivers</article-title>. <source>Biol. Conserv.</source> <volume>232</volume>, <fpage>8</fpage>&#x2013;<lpage>27</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.biocon.2019.01.020</pub-id></mixed-citation></ref>
<ref id="ref122"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Santos-Medell&#x00ED;n</surname><given-names>C.</given-names></name> <name><surname>Edwards</surname><given-names>J.</given-names></name> <name><surname>Liechty</surname><given-names>Z.</given-names></name> <name><surname>Nguyen</surname><given-names>B.</given-names></name> <name><surname>Sundaresan</surname><given-names>V.</given-names></name></person-group> (<year>2017</year>). <article-title>Drought stress results in a compartment-specific restructuring of the rice root-associated microbiomes</article-title>. <source>MBio</source> <volume>8</volume>:<fpage>e01011&#x2013;17</fpage>. doi: <pub-id pub-id-type="doi">10.1128/mbio.00764-17</pub-id>, <pub-id pub-id-type="pmid">28720730</pub-id></mixed-citation></ref>
<ref id="ref123"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Schulz</surname><given-names>A. N.</given-names></name> <name><surname>Lucardi</surname><given-names>R. D.</given-names></name> <name><surname>Marsico</surname><given-names>T. D.</given-names></name></person-group> (<year>2019</year>). <article-title>Successful invasions and failed biocontrol: the role of antagonistic species interactions</article-title>. <source>Bioscience</source> <volume>69</volume>, <fpage>711</fpage>&#x2013;<lpage>724</lpage>. doi: <pub-id pub-id-type="doi">10.1093/biosci/biz075</pub-id></mixed-citation></ref>
<ref id="ref124"><mixed-citation publication-type="book"><person-group person-group-type="author"><name><surname>Sharma</surname><given-names>S.</given-names></name> <name><surname>Chandra</surname><given-names>D.</given-names></name> <name><surname>Sharma</surname><given-names>A. K.</given-names></name></person-group> (<year>2020</year>). &#x201C;<chapter-title>Rhizosphere plant&#x2013;microbe interactions under abiotic stress</chapter-title>&#x201D; in <source>Rhizosphere Biology: Interactions between Microbes and Plants</source>. eds. <person-group person-group-type="editor"><name><surname>Vadakattu</surname><given-names>V. S. R. G.</given-names></name> <name><surname>Anil</surname><given-names>K. S.</given-names></name></person-group> (<publisher-loc>Singapore</publisher-loc>: <publisher-name>Springer</publisher-name>), <fpage>195</fpage>&#x2013;<lpage>216</lpage>.</mixed-citation></ref>
<ref id="ref125"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Sheoran</surname><given-names>A. R.</given-names></name> <name><surname>Lakra</surname><given-names>N.</given-names></name> <name><surname>Saharan</surname><given-names>B. S.</given-names></name> <name><surname>Luhach</surname><given-names>A.</given-names></name> <name><surname>Kumar</surname><given-names>R.</given-names></name> <name><surname>Seth</surname><given-names>C. S.</given-names></name> <etal/></person-group>. (<year>2025</year>). <article-title>Enhancing plant disease resistance: insights from biocontrol agent strategies</article-title>. <source>J. Plant Growth Regul.</source> <volume>44</volume>, <fpage>436</fpage>&#x2013;<lpage>459</lpage>. doi: <pub-id pub-id-type="doi">10.1007/s00344-024-11480-y</pub-id></mixed-citation></ref>
<ref id="ref126"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Singh</surname><given-names>B. K.</given-names></name> <name><surname>Delgado-Baquerizo</surname><given-names>M.</given-names></name> <name><surname>Egidi</surname><given-names>E.</given-names></name> <name><surname>Guirado</surname><given-names>E.</given-names></name> <name><surname>Leach</surname><given-names>J. E.</given-names></name> <name><surname>Liu</surname><given-names>H.</given-names></name> <etal/></person-group>. (<year>2023</year>). <article-title>Climate change impacts on plant pathogens, food security and paths forward</article-title>. <source>Nat. Rev. Microbiol.</source> <volume>21</volume>, <fpage>640</fpage>&#x2013;<lpage>656</lpage>. doi: <pub-id pub-id-type="doi">10.1038/s41579-023-00900-7</pub-id>, <pub-id pub-id-type="pmid">37131070</pub-id></mixed-citation></ref>
<ref id="ref127"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Sorahinobar</surname><given-names>M.</given-names></name> <name><surname>Yusefieh</surname><given-names>N.</given-names></name> <name><surname>Rezayian</surname><given-names>M.</given-names></name> <name><surname>Shahbazi</surname><given-names>S.</given-names></name></person-group> (<year>2025</year>). <article-title>Multifaceted role of <italic>Trichoderma harzianum</italic> isolates in mitigating drought stress and promoting adaptive responses in barley cultivars</article-title>. <source>Sci. Rep.</source> <volume>15</volume>:<fpage>26552</fpage>. doi: <pub-id pub-id-type="doi">10.1038/s41598-025-08922-2</pub-id>, <pub-id pub-id-type="pmid">40695871</pub-id></mixed-citation></ref>
<ref id="ref128"><mixed-citation publication-type="book"><person-group person-group-type="author"><name><surname>Srivastava</surname><given-names>Y.</given-names></name></person-group> (<year>2019</year>). &#x201C;<chapter-title>Climate change: a challenge for postharvest management, food loss, food quality, and food security</chapter-title>&#x201D; in <source>Climate Change and Agricultural Ecosystems: Current Challenges and Adaptation</source>. eds. <person-group person-group-type="editor"><name><surname>Choudhary</surname><given-names>K. K.</given-names></name> <name><surname>Kumar</surname><given-names>A.</given-names></name> <name><surname>Singh</surname><given-names>A. K.</given-names></name></person-group> (<publisher-name>Woodhead Publishing, Elsevier Inc.</publisher-name>), <fpage>355</fpage>&#x2013;<lpage>377</lpage>. doi: <pub-id pub-id-type="doi">10.1016/C2018-0-00420-X</pub-id></mixed-citation></ref>
<ref id="ref129"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Stenberg</surname><given-names>J. A.</given-names></name></person-group> (<year>2017</year>). <article-title>A conceptual framework for integrated pest management</article-title>. <source>Trends Plant Sci.</source> <volume>22</volume>, <fpage>759</fpage>&#x2013;<lpage>769</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.tplants.2017.06.010</pub-id>, <pub-id pub-id-type="pmid">28687452</pub-id></mixed-citation></ref>
<ref id="ref130"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Su</surname><given-names>L.</given-names></name> <name><surname>Zhang</surname><given-names>L.</given-names></name> <name><surname>Nie</surname><given-names>D.</given-names></name> <name><surname>Kuramae</surname><given-names>E. E.</given-names></name> <name><surname>Shen</surname><given-names>B.</given-names></name> <name><surname>Shen</surname><given-names>Q.</given-names></name></person-group> (<year>2020</year>). <article-title>Bacterial tomato pathogen <italic>Ralstonia solanacearum</italic> invasion modulates rhizosphere compounds and facilitates the cascade effect of fungal pathogen <italic>Fusarium solani</italic></article-title>. <source>Microorganisms</source> <volume>8</volume>:<fpage>806</fpage>. doi: <pub-id pub-id-type="doi">10.3390/microorganisms8060806</pub-id>, <pub-id pub-id-type="pmid">32471167</pub-id></mixed-citation></ref>
<ref id="ref131"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Suman</surname><given-names>J.</given-names></name> <name><surname>Rakshit</surname><given-names>A.</given-names></name> <name><surname>Ogireddy</surname><given-names>S. D.</given-names></name> <name><surname>Singh</surname><given-names>S.</given-names></name> <name><surname>Gupta</surname><given-names>C.</given-names></name> <name><surname>Chandrakala</surname><given-names>J.</given-names></name></person-group> (<year>2022</year>). <article-title>Microbiome as a key player in sustainable agriculture and human health</article-title>. <source>Front. Soil Sci.</source> <volume>2</volume>:<fpage>821589</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fsoil.2022.821589</pub-id></mixed-citation></ref>
<ref id="ref132"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Sun</surname><given-names>Y.</given-names></name> <name><surname>Guo</surname><given-names>J.</given-names></name> <name><surname>Mur</surname><given-names>L. A. J.</given-names></name> <name><surname>Xu</surname><given-names>X.</given-names></name> <name><surname>Chen</surname><given-names>H.</given-names></name> <name><surname>Yang</surname><given-names>Y.</given-names></name> <etal/></person-group>. (<year>2024</year>). <article-title>Nitrogen starvation modulates the sensitivity of rhizobacterial community to drought stress in <italic>Stevia rebaudiana</italic></article-title>. <source>J. Environ. Manag.</source> <volume>354</volume>:<fpage>120486</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.jenvman.2024.120486</pub-id></mixed-citation></ref>
<ref id="ref133"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Sun</surname><given-names>T.</given-names></name> <name><surname>Liu</surname><given-names>H.</given-names></name> <name><surname>Wang</surname><given-names>N.</given-names></name> <name><surname>Huang</surname><given-names>M.</given-names></name> <name><surname>Banerjee</surname><given-names>S.</given-names></name> <name><surname>Jousset</surname><given-names>A.</given-names></name> <etal/></person-group>. (<year>2025</year>). <article-title>Interactions with native microbial keystone taxa enhance the biocontrol efficiency of <italic>Streptomyces</italic></article-title>. <source>Microbiome</source> <volume>13</volume>:<fpage>126</fpage>. doi: <pub-id pub-id-type="doi">10.1186/s40168-025-02120-y</pub-id>, <pub-id pub-id-type="pmid">40390122</pub-id></mixed-citation></ref>
<ref id="ref134"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Suzuki</surname><given-names>N.</given-names></name> <name><surname>Rivero</surname><given-names>R. M.</given-names></name> <name><surname>Shulaev</surname><given-names>V.</given-names></name> <name><surname>Blumwald</surname><given-names>E.</given-names></name> <name><surname>Mittler</surname><given-names>R.</given-names></name></person-group> (<year>2014</year>). <article-title>Abiotic and biotic stress combinations</article-title>. <source>New Phytol.</source> <volume>203</volume>, <fpage>32</fpage>&#x2013;<lpage>43</lpage>. doi: <pub-id pub-id-type="doi">10.1111/nph.12797</pub-id>, <pub-id pub-id-type="pmid">24720847</pub-id></mixed-citation></ref>
<ref id="ref135"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Tang</surname><given-names>Z.</given-names></name> <name><surname>Tan</surname><given-names>W.</given-names></name> <name><surname>Li</surname><given-names>R.</given-names></name> <name><surname>Weng</surname><given-names>L.</given-names></name> <name><surname>Chen</surname><given-names>X.</given-names></name> <name><surname>Xi</surname><given-names>B.</given-names></name> <etal/></person-group>. (<year>2025</year>). <article-title>Advances in rhizosphere microbiome and rhizosphere immunity effect: a review</article-title>. <source>J. Agric. Food Chem.</source> <volume>72</volume>, <fpage>14707</fpage>&#x2013;<lpage>14721</lpage>. doi: <pub-id pub-id-type="doi">10.1021/acs.jafc.5c02003</pub-id></mixed-citation></ref>
<ref id="ref136"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Thangaraj</surname><given-names>P.</given-names></name> <name><surname>Balamurali</surname><given-names>A. S.</given-names></name> <name><surname>Muthusamy</surname><given-names>N.</given-names></name></person-group> (<year>2025</year>). <article-title>Biological control of <italic>Trichoderma</italic> spp.: mechanisms of action against phytopathogens, insect pests, and its multifaceted roles in agro-ecosystems</article-title>. <source>Environ. Conserv. J.</source> <volume>26</volume>, <fpage>302</fpage>&#x2013;<lpage>314</lpage>. doi: <pub-id pub-id-type="doi">10.36953/ecj.28922909</pub-id></mixed-citation></ref>
<ref id="ref137"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Thepbandit</surname><given-names>W.</given-names></name> <name><surname>Athinuwat</surname><given-names>D.</given-names></name></person-group> (<year>2024</year>). <article-title>Rhizosphere microorganisms supply availability of soil nutrients and induce plant defense</article-title>. <source>Microorganisms</source> <volume>12</volume>:<fpage>558</fpage>. doi: <pub-id pub-id-type="doi">10.3390/microorganisms12030558</pub-id>, <pub-id pub-id-type="pmid">38543610</pub-id></mixed-citation></ref>
<ref id="ref138"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Trabelsi</surname><given-names>D.</given-names></name> <name><surname>Mhamdi</surname><given-names>R.</given-names></name></person-group> (<year>2013</year>). <article-title>Microbial inoculants and their impact on soil microbial communities: a review</article-title>. <source>Biomed. Res. Int.</source> <volume>2013</volume>:<fpage>863240</fpage>. doi: <pub-id pub-id-type="doi">10.1155/2013/863240</pub-id>, <pub-id pub-id-type="pmid">23957006</pub-id></mixed-citation></ref>
<ref id="ref139"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Trivedi</surname><given-names>P.</given-names></name> <name><surname>Batista</surname><given-names>B. D.</given-names></name> <name><surname>Bazany</surname><given-names>K. E.</given-names></name> <name><surname>Singh</surname><given-names>B. K.</given-names></name></person-group> (<year>2022</year>). <article-title>Plant&#x2013;microbiome interactions under a changing world: responses, consequences and perspectives</article-title>. <source>New Phytol.</source> <volume>234</volume>, <fpage>1951</fpage>&#x2013;<lpage>1959</lpage>. doi: <pub-id pub-id-type="doi">10.1111/nph.18016</pub-id>, <pub-id pub-id-type="pmid">35118660</pub-id></mixed-citation></ref>
<ref id="ref140"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Tyagi</surname><given-names>J.</given-names></name> <name><surname>Mishra</surname><given-names>A.</given-names></name> <name><surname>Kumari</surname><given-names>S.</given-names></name> <name><surname>Singh</surname><given-names>S.</given-names></name> <name><surname>Agarwal</surname><given-names>H.</given-names></name> <name><surname>Pudake</surname><given-names>R. N.</given-names></name> <etal/></person-group>. (<year>2023</year>). <article-title>Deploying a microbial consortium of <italic>Serendipita indica</italic>, <italic>Rhizophagus intraradices</italic>, and <italic>Azotobacter chroococcum</italic> to boost drought tolerance in maize</article-title>. <source>Environ. Exp. Bot.</source> <volume>206</volume>:<fpage>105142</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.envexpbot.2022.105142</pub-id></mixed-citation></ref>
<ref id="ref141"><mixed-citation publication-type="other"><collab id="coll1">United Nations</collab> (<year>2019</year>). 9.7 billion on earth by 2050, but growth rate slowing, says new UN population report. United Nations. Available online at: <ext-link xlink:href="https://www.un.org/en/academic-impact/97-billion-earth-2050-growth-rate-slowing-says-new-un-population-report" ext-link-type="uri">https://www.un.org/en/academic-impact/97-billion-earth-2050-growth-rate-slowing-says-new-un-population-report</ext-link> (Accessed September 12, 2025).</mixed-citation></ref>
<ref id="ref142"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Van Dijk</surname><given-names>M.</given-names></name> <name><surname>Morley</surname><given-names>T.</given-names></name> <name><surname>Rau</surname><given-names>M. L.</given-names></name> <name><surname>Saghai</surname><given-names>Y.</given-names></name></person-group> (<year>2021</year>). <article-title>A meta-analysis of projected global food demand and population at risk of hunger for the period 2010&#x2013;2050</article-title>. <source>Nat. Food</source> <volume>2</volume>, <fpage>494</fpage>&#x2013;<lpage>501</lpage>. doi: <pub-id pub-id-type="doi">10.1038/s43016-021-00322-9</pub-id>, <pub-id pub-id-type="pmid">37117684</pub-id></mixed-citation></ref>
<ref id="ref143"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Vishwakarma</surname><given-names>K.</given-names></name> <name><surname>Kumar</surname><given-names>N.</given-names></name> <name><surname>Shandilya</surname><given-names>C.</given-names></name> <name><surname>Mohapatra</surname><given-names>S.</given-names></name> <name><surname>Bhayana</surname><given-names>S.</given-names></name> <name><surname>Varma</surname><given-names>A.</given-names></name></person-group> (<year>2020</year>). <article-title>Revisiting plant&#x2013;microbe interactions and microbial consortia application for enhancing sustainable agriculture: a review</article-title>. <source>Front. Microbiol.</source> <volume>11</volume>:<fpage>560406</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fmicb.2020.560406</pub-id>, <pub-id pub-id-type="pmid">33408698</pub-id></mixed-citation></ref>
<ref id="ref144"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Wahab</surname><given-names>A.</given-names></name> <name><surname>Muhammad</surname><given-names>M.</given-names></name> <name><surname>Munir</surname><given-names>A.</given-names></name> <name><surname>Abdi</surname><given-names>G.</given-names></name> <name><surname>Zaman</surname><given-names>W.</given-names></name> <name><surname>Ayaz</surname><given-names>A.</given-names></name> <etal/></person-group>. (<year>2023</year>). <article-title>Role of arbuscular mycorrhizal fungi in regulating growth, enhancing productivity, and potentially influencing ecosystems under abiotic and biotic stresses</article-title>. <source>Plants</source> <volume>12</volume>:<fpage>3102</fpage>. doi: <pub-id pub-id-type="doi">10.3390/plants12173102</pub-id>, <pub-id pub-id-type="pmid">37687353</pub-id></mixed-citation></ref>
<ref id="ref145"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Wan</surname><given-names>T.</given-names></name> <name><surname>Zhao</surname><given-names>H.</given-names></name> <name><surname>Wang</surname><given-names>W.</given-names></name></person-group> (<year>2017</year>). <article-title>Effect of biocontrol agent <italic>Bacillus amyloliquefaciens</italic> SN16-1 and plant pathogen <italic>fusarium oxysporum</italic> on tomato rhizosphere bacterial community composition</article-title>. <source>Biol. Control</source> <volume>112</volume>, <fpage>1</fpage>&#x2013;<lpage>9</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.biocontrol.2017.05.014</pub-id></mixed-citation></ref>
<ref id="ref146"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Whipps</surname><given-names>J. M.</given-names></name></person-group> (<year>2001</year>). <article-title>Microbial interactions and biocontrol in the rhizosphere</article-title>. <source>J. Exp. Bot.</source> <volume>52</volume>, <fpage>487</fpage>&#x2013;<lpage>511</lpage>. doi: <pub-id pub-id-type="doi">10.1093/jexbot/52.suppl_1.487</pub-id></mixed-citation></ref>
<ref id="ref147"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Wilson</surname><given-names>K.</given-names></name> <name><surname>Arunachalam</surname><given-names>S.</given-names></name></person-group> (<year>2024</year>). <article-title>Microbiome transition mediated plant immune response to <italic>Alternaria solani</italic> (Ellis &#x0026; Martin) Jones &#x0026; Grout infection in tomato (<italic>Solanum lycopersicum</italic> L.)</article-title>. <source>Heliyon</source> <volume>10</volume>:<fpage>e37203</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.heliyon.2024.e37203</pub-id>, <pub-id pub-id-type="pmid">39296181</pub-id></mixed-citation></ref>
<ref id="ref148"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Woo</surname><given-names>S. L.</given-names></name> <name><surname>Pepe</surname><given-names>O.</given-names></name></person-group> (<year>2018</year>). <article-title>Microbial consortia: promising probiotics as plant biostimulants for sustainable agriculture</article-title>. <source>Front. Plant Sci.</source> <volume>9</volume>:<fpage>1801</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fpls.2018.01801</pub-id>, <pub-id pub-id-type="pmid">30564264</pub-id></mixed-citation></ref>
<ref id="ref149"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Yousaf</surname><given-names>M. I.</given-names></name> <name><surname>Riaz</surname><given-names>M. W.</given-names></name> <name><surname>Shehzad</surname><given-names>A.</given-names></name> <name><surname>Jamil</surname><given-names>S.</given-names></name> <name><surname>Shahzad</surname><given-names>R.</given-names></name> <name><surname>Kanwal</surname><given-names>S.</given-names></name> <etal/></person-group>. (<year>2023</year>). <article-title>Responses of maize hybrids to water stress conditions at different developmental stages: accumulation of reactive oxygen species, activity of enzymatic antioxidants and degradation in kernel quality traits</article-title>. <source>PeerJ</source> <volume>11</volume>:<fpage>e14983</fpage>. doi: <pub-id pub-id-type="doi">10.7717/peerj.14983</pub-id>, <pub-id pub-id-type="pmid">36967996</pub-id></mixed-citation></ref>
<ref id="ref150"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Zahra</surname><given-names>N.</given-names></name> <name><surname>Hafeez</surname><given-names>M. B.</given-names></name> <name><surname>Al Shukaily</surname><given-names>M.</given-names></name> <name><surname>Al-Sadi</surname><given-names>A. M.</given-names></name> <name><surname>Siddique</surname><given-names>K. H.</given-names></name> <name><surname>Farooq</surname><given-names>M.</given-names></name></person-group> (<year>2023</year>). <article-title>Influence of abiotic stresses on disease infestation in plants</article-title>. <source>Physiol. Mol. Plant Pathol.</source> <volume>127</volume>:<fpage>102125</fpage>. doi: <pub-id pub-id-type="doi">10.1016/j.pmpp.2023.102125</pub-id></mixed-citation></ref>
<ref id="ref151"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Zandalinas</surname><given-names>S. I.</given-names></name> <name><surname>Balfag&#x00F3;n</surname><given-names>D.</given-names></name> <name><surname>G&#x00F3;mez-Cadenas</surname><given-names>A.</given-names></name> <name><surname>Mittler</surname><given-names>R.</given-names></name></person-group> (<year>2022</year>). <article-title>Plant responses to climate change: metabolic changes under combined abiotic stresses</article-title>. <source>J. Exp. Bot.</source> <volume>73</volume>, <fpage>3339</fpage>&#x2013;<lpage>3354</lpage>. doi: <pub-id pub-id-type="doi">10.1093/jxb/erac073</pub-id>, <pub-id pub-id-type="pmid">35192700</pub-id></mixed-citation></ref>
<ref id="ref152"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Zayadan</surname><given-names>B. K.</given-names></name> <name><surname>Matorin</surname><given-names>D. N.</given-names></name> <name><surname>Baimakhanova</surname><given-names>G. B.</given-names></name> <name><surname>Bolathan</surname><given-names>K.</given-names></name> <name><surname>Oraz</surname><given-names>G. D.</given-names></name> <name><surname>Sadanov</surname><given-names>A. K.</given-names></name></person-group> (<year>2014</year>). <article-title>Promising microbial consortia for producing biofertilizers for rice fields</article-title>. <source>Microbiology</source> <volume>83</volume>, <fpage>391</fpage>&#x2013;<lpage>397</lpage>. doi: <pub-id pub-id-type="doi">10.1134/s0026261714040171</pub-id></mixed-citation></ref>
<ref id="ref153"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Zhang</surname><given-names>N.</given-names></name> <name><surname>Wang</surname><given-names>Z.</given-names></name> <name><surname>Shao</surname><given-names>J.</given-names></name> <name><surname>Xu</surname><given-names>Z.</given-names></name> <name><surname>Liu</surname><given-names>Y.</given-names></name> <name><surname>Xun</surname><given-names>W.</given-names></name> <etal/></person-group>. (<year>2023</year>). <article-title>Biocontrol mechanisms of <italic>Bacillus</italic>: improving the efficiency of green agriculture</article-title>. <source>Microb. Biotechnol.</source> <volume>16</volume>, <fpage>2250</fpage>&#x2013;<lpage>2263</lpage>. doi: <pub-id pub-id-type="doi">10.1111/1751-7915.14348</pub-id>, <pub-id pub-id-type="pmid">37837627</pub-id></mixed-citation></ref>
<ref id="ref154"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Zhang</surname><given-names>Y.</given-names></name> <name><surname>Xiao</surname><given-names>J.</given-names></name> <name><surname>Yang</surname><given-names>K.</given-names></name> <name><surname>Wang</surname><given-names>Y.</given-names></name> <name><surname>Tian</surname><given-names>Y.</given-names></name> <name><surname>Liang</surname><given-names>Z.</given-names></name></person-group> (<year>2022</year>). <article-title>Transcriptomic and metabonomic insights into the biocontrol mechanism of <italic>Trichoderma asperellum</italic> M45a against watermelon fusarium wilt</article-title>. <source>PLoS One</source> <volume>17</volume>:<fpage>e0272702</fpage>. doi: <pub-id pub-id-type="doi">10.1371/journal.pone.0272702</pub-id>, <pub-id pub-id-type="pmid">35947630</pub-id></mixed-citation></ref>
<ref id="ref155"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Zheng</surname><given-names>W.</given-names></name> <name><surname>Zeng</surname><given-names>S.</given-names></name> <name><surname>Bais</surname><given-names>H.</given-names></name> <name><surname>LaManna</surname><given-names>J. M.</given-names></name> <name><surname>Hussey</surname><given-names>D. S.</given-names></name> <name><surname>Jacobson</surname><given-names>D. L.</given-names></name> <etal/></person-group>. (<year>2018</year>). <article-title>Plant growth-promoting rhizobacteria (PGPR) reduce evaporation and increase soil water retention</article-title>. <source>Water Resour. Res.</source> <volume>54</volume>, <fpage>3673</fpage>&#x2013;<lpage>3687</lpage>. doi: <pub-id pub-id-type="doi">10.1029/2018wr022656</pub-id></mixed-citation></ref>
<ref id="ref156"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Zhu</surname><given-names>L.</given-names></name> <name><surname>Huang</surname><given-names>J.</given-names></name> <name><surname>Lu</surname><given-names>X.</given-names></name> <name><surname>Zhou</surname><given-names>C.</given-names></name></person-group> (<year>2022</year>). <article-title>Development of plant systemic resistance by beneficial rhizobacteria: recognition, initiation, elicitation and regulation</article-title>. <source>Front. Plant Sci.</source> <volume>13</volume>:<fpage>952397</fpage>. doi: <pub-id pub-id-type="doi">10.3389/fpls.2022.952397</pub-id>, <pub-id pub-id-type="pmid">36017257</pub-id></mixed-citation></ref>
</ref-list>
<fn-group>
<fn fn-type="custom" custom-type="edited-by" id="fn0001">
<p>Edited by: <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/1310841/overview">Ahmad Ali</ext-link>, Florida Agricultural and Mechanical University, United States</p>
</fn>
<fn fn-type="custom" custom-type="reviewed-by" id="fn0002">
<p>Reviewed by: <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/2168286/overview">Junwei Peng</ext-link>, Chinese Academy of Sciences (CAS), China</p>
<p><ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/3227122/overview">Dipayan Das</ext-link>, Royal Global University, India</p>
</fn>
</fn-group>
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