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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Earth Sci.</journal-id>
<journal-title-group>
<journal-title>Frontiers in Earth Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Earth Sci.</abbrev-journal-title>
</journal-title-group>
<issn pub-type="epub">2296-6463</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
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<article-meta>
<article-id pub-id-type="publisher-id">1752600</article-id>
<article-id pub-id-type="doi">10.3389/feart.2026.1752600</article-id>
<article-version article-version-type="Version of Record" vocab="NISO-RP-8-2008"/>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Systematic Review</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>Microplastic pollution in aquatic environments: a systematic review of bacterial degradation efficacy, mechanisms, and future pathways</article-title>
<alt-title alt-title-type="left-running-head">Barwant et al.</alt-title>
<alt-title alt-title-type="right-running-head">
<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/feart.2026.1752600">10.3389/feart.2026.1752600</ext-link>
</alt-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Barwant</surname>
<given-names>Mukul Machhindra</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="corresp" rid="c001">&#x2a;</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2097125"/>
<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 - original draft</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; review &#x26; editing" vocab-term-identifier="https://credit.niso.org/contributor-roles/Writing - review &#x26; editing/">Writing - review and editing</role>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Ali</surname>
<given-names>Usman Mohammed</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/3062255"/>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; review &#x26; editing" vocab-term-identifier="https://credit.niso.org/contributor-roles/Writing - review &#x26; editing/">Writing - review and editing</role>
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</contrib>
<contrib contrib-type="author">
<name>
<surname>Tolani</surname>
<given-names>Terefu Regassa</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; review &#x26; editing" vocab-term-identifier="https://credit.niso.org/contributor-roles/Writing - review &#x26; editing/">Writing - review and editing</role>
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<aff id="aff1">
<label>1</label>
<institution>Department of Botany, Sanjivani Arts Commerce and Science College Kopargoan</institution>, <city>Ahmednagar</city>, <state>Maharashtra</state>, <country country="IN">India</country>
</aff>
<aff id="aff2">
<label>2</label>
<institution>Department of Plant Sciences, Faculty of Agriculture, Wollega University</institution>, <city>Shambu</city>, <country country="ET">Ethiopia</country>
</aff>
<author-notes>
<corresp id="c001">
<label>&#x2a;</label>Correspondence: Mukul Machhindra Barwant, <email xlink:href="mailto:mukulbarwant97@gmail.com">mukulbarwant97@gmail.com</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>14</volume>
<elocation-id>1752600</elocation-id>
<history>
<date date-type="received">
<day>23</day>
<month>11</month>
<year>2025</year>
</date>
<date date-type="rev-recd">
<day>13</day>
<month>01</month>
<year>2026</year>
</date>
<date date-type="accepted">
<day>16</day>
<month>01</month>
<year>2026</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2026 Barwant, Ali and Tolani.</copyright-statement>
<copyright-year>2026</copyright-year>
<copyright-holder>Barwant, Ali and Tolani</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 pervasive accumulation of microplastics (MPs) in aquatic ecosystems constitutes a critical environmental threat, necessitating sustainable remediation strategies. Bacterial degradation has emerged as a promising solution, yet a systematic synthesis of its efficacy, mechanisms, and practical feasibility is lacking. Following PRISMA guidelines, a comprehensive search of Scopus, Web of Science, and PubMed databases (2000&#x2013;2025) was conducted, yielding 80 eligible studies out of 639 identified records. Qualitative thematic synthesis was employed to analyze the evidence. The analysis reveals a conserved consortium of bacteria primarily <italic>Pseudomonas</italic>, <italic>Bacillus</italic>, and <italic>Rhodococcus</italic> capable of colonizing the &#x201c;plastisphere&#x201d; and degrading major polymers through specific enzymatic pathways. Hydrolases (e.g., PETase) enable rapid depolymerization of hydrolysable polymers like PET, while oxidoreductases (e.g., alkane hydroxylases) slowly oxidize recalcitrant polyolefins (PE, PP). Degradation efficacy is highly polymer-dependent, with PET showing the most promise (up to 50% mass loss in days) compared to significantly slower PE/PP degradation (&#x3c;12% over months). Key factors influencing kinetics include temperature, pH, nutrient availability, and the synergistic effects of microbial consortia within biofilms. However, translation to field applications faces formidable barriers, including ecological competition, scalability challenges, and difficulties in monitoring efficacy in open environments. Critical knowledge gaps persist regarding long-term environmental fate, ecotoxicity of degradation by-products, and the rational design of effective microbial consortia. While bacterial degradation presents a scientifically validated mechanism for MP bioremediation, its near-term application is most viable in controlled, <italic>ex-situ</italic> systems like enzymatic recycling of PET. For <italic>in-situ</italic> remediation, a tailored, hybrid bioaugmentation-biosimulation approach is recommended, though significant research and policy hurdles remain. Future work must prioritize standardized methods, long-term ecological studies, and integrated risk assessment to translate this promising biotechnology into practical environmental solutions.</p>
</abstract>
<abstract abstract-type="graphical">
<title>Graphical Abstract</title>
<p>
<fig>
<graphic xlink:href="FEART_feart-2026-1752600_wc_abs.tif" position="anchor">
<alt-text content-type="machine-generated">Diagram illustrating bacterial degradation of aquatic microplastics. It shows sources of microplastic pollution in water. Bacterial colonization involves species like Pseudomonas and Bacillus. Rapid hydrolysis, using hydrolases such as PETase, results in high efficacy with fifty percent mass loss in days. Slow oxidation involves oxidoreductases with low efficacy under twelve percent mass loss in months. Barriers include competition and scalability, suggesting near-term enzymatic recycling and future recommendations for hybrid bioaugmentation.</alt-text>
</graphic>
</fig>
</p>
</abstract>
<kwd-group>
<kwd>bacterial consortia</kwd>
<kwd>bioremediation</kwd>
<kwd>circular economy</kwd>
<kwd>enzymatic depolymerization</kwd>
<kwd>microplastic biodegradation</kwd>
<kwd>plastisphere</kwd>
<kwd>polymer persistence</kwd>
</kwd-group>
<funding-group>
<funding-statement>The author(s) declared that financial support was not received for this work and/or its publication.</funding-statement>
</funding-group>
<counts>
<fig-count count="7"/>
<table-count count="9"/>
<equation-count count="0"/>
<ref-count count="80"/>
<page-count count="00"/>
</counts>
<custom-meta-group>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Sedimentology, Stratigraphy and Diagenesis</meta-value>
</custom-meta>
</custom-meta-group>
</article-meta>
</front>
<body>
<sec sec-type="intro" id="s1">
<label>1</label>
<title>Introduction</title>
<sec id="s1-1">
<label>1.1</label>
<title>The global challenge of microplastic pollution in aquatic ecosystems</title>
<p>The ubiquity of plastic pollution represents one of the most pressing environmental challenges of the Anthropocene. Global plastic production has surged to exceed 400 million metric tons annually, a substantial portion of which enters and accumulates in natural ecosystems (<xref ref-type="bibr" rid="B56">Rangel-Buitrago and Galgani, 2026</xref>). Through processes of mechanical abrasion, photodegradation, and biological weathering, larger plastic debris undergoes fragmentation into microplastics (MPs), defined as synthetic polymer particles less than 5 mm in diameter (<xref ref-type="bibr" rid="B39">Matavos-Aramyan, 2024</xref>; <xref ref-type="bibr" rid="B23">Jolaosho et al., 2025</xref>). These micropollutants have permeated every corner of the planet&#x2019;s hydrosphere, from pristine alpine lakes and deep-sea trenches to urban rivers and coastal estuaries (<xref ref-type="bibr" rid="B69">Tarigan et al., 2025</xref>). Their small size, combined with their persistent nature and high surface-area-to-volume ratio, renders them bioavailable to a vast range of aquatic organisms, facilitating trophic transfer and posing a significant threat to biodiversity, ecosystem function, and potentially human health through seafood consumption (<xref ref-type="bibr" rid="B8">Bulannga and Schmidt, 2024</xref>; <xref ref-type="bibr" rid="B49">Pal et al., 2025</xref>). The environmental persistence of conventional polymers, which may span centuries, underscores the urgency of developing effective remediation strategies (<xref ref-type="bibr" rid="B5">Beena Unni and Muringayil Joseph, 2024</xref>).</p>
</sec>
<sec id="s1-2">
<label>1.2</label>
<title>Sources, pathways, and ecological impacts of microplastics</title>
<p>Microplastics originate from a complex mixture of primary sources, such as microbeads from personal care products and pre-production pellets, and secondary sources derived from the breakdown of larger items like packaging materials and fishing gear (<xref ref-type="bibr" rid="B2">Ashrafy et al., 2023</xref>; <xref ref-type="bibr" rid="B46">Nkin, 2025</xref>). Pathways into aquatic environments are multifaceted, including wastewater discharge, agricultural runoff, atmospheric deposition, and improper waste management (<xref ref-type="bibr" rid="B59">Rothman et al., 2023</xref>; <xref ref-type="bibr" rid="B39">Matavos-Aramyan, 2024</xref>). Once introduced, their ecological impacts are multifaceted. Physically, ingestion can lead to gut blockage, false satiation, and internal injury in aquatic fauna (<xref ref-type="bibr" rid="B35">Liu and Li, 2025</xref>). Chemically, MPs can act as vectors for toxic additives (e.g., plasticizers, flame retardants) and hydrophobic environmental contaminants (e.g., heavy metals, persistent organic pollutants), concentrating them and enhancing their bioavailability (<xref ref-type="bibr" rid="B41">Mbachu et al., 2020</xref>; <xref ref-type="bibr" rid="B57">Rashid et al., 2025</xref>. Furthermore, the plastisphere the unique microbial community colonizing MP surfaces can include potential pathogens and facilitate the transport of antibiotic resistance genes, adding a complex biological dimension to the threat (<xref ref-type="bibr" rid="B72">Wu et al., 2024</xref>).</p>
</sec>
<sec id="s1-3">
<label>1.3</label>
<title>Bioremediation as a sustainable mitigation strategy: the role of bacteria</title>
<p>Conventional methods for MP removal from water bodies, such as filtration and coagulation, are often energy-intensive, costly, and impractical for large-scale application in open environments. They primarily focus on sequestration rather than degradation, leading to a secondary waste problem (<xref ref-type="bibr" rid="B55">Puteri et al., 2025</xref>). In this context, bioremediation the use of biological organisms to detoxify or remove pollutants has emerged as a promising, eco-friendly, and potentially scalable alternative. Among biological agents, bacteria are at the forefront of MP bio-remediation research due to their ubiquitous distribution, metabolic versatility, and rapid reproduction rates (<xref ref-type="bibr" rid="B33">Kuppan et al., 2024</xref>). A growing body of evidence has identified diverse bacterial taxa, including genera such as <italic>Pseudomonas</italic>, <italic>Bacillus</italic>, <italic>Rhodococcus</italic>, and <italic>Ideonella</italic>, capable of adhering to plastic surfaces and utilizing them as a carbon and energy source through the secretion of specific extracellular enzymes like hydrolases and laccases (<xref ref-type="bibr" rid="B74">Yadav et al., 2025</xref>). This enzymatic activity can break down the long-chain polymers into smaller oligomers, monomers, and ultimately, carbon dioxide and water, offering a pathway for the complete mineralization of plastic waste.</p>
</sec>
<sec id="s1-4">
<label>1.4</label>
<title>Rationale and knowledge gaps justifying a systematic review</title>
<p>Despite a surge in primary research articles over the past decade, the field of bacterial degradation of microplastics remains characterized by significant fragmentation and heterogeneity. Studies are often conducted in isolation, using disparate methodologies, different bacterial strains, and a wide array of polymer types, making cross-comparison and the drawing of general conclusions challenging (<xref ref-type="bibr" rid="B54">Przygoda-Ku&#x15b; et al., 2025</xref>). Critical knowledge gaps persist regarding the specific enzymatic mechanisms and their genetic regulation, the kinetics of degradation under environmentally relevant conditions (as opposed to optimized lab settings), the efficacy of bacterial consortia versus individual isolates, and the potential ecotoxicological risks associated with degradation by-products (<xref ref-type="bibr" rid="B51">Parab et al., 2025</xref>). While several narrative reviews have summarized progress in this area, a comprehensive, methodologically rigorous, and transparent synthesis of the global evidence is lacking. A systematic review is therefore imperative to consolidate the scattered findings, critically appraise the quality of the evidence, quantify the overall efficacy where possible, and identify the most robust and promising pathways for future research and application.</p>
</sec>
<sec id="s1-5">
<label>1.5</label>
<title>Review objectives and research questions</title>
<p>This systematic review aims to provide a comprehensive and critical synthesis of the current state of knowledge on the bacterial degradation of microplastics in aquatic environments. Guided by the PRISMA (Preferred Reporting Items for Systematic Reviews and Meta-Analyses) guidelines, this review seeks to address the following specific research questions:<list list-type="order">
<list-item>
<p>What is the taxonomic and functional diversity of bacteria identified as capable of degrading the most prevalent microplastic polymers (e.g., polyethylene, polypropylene, polystyrene, PET) in aquatic environments?</p>
</list-item>
<list-item>
<p>What are the primary enzymatic mechanisms and metabolic pathways involved in the bacterial degradation of these microplastics?</p>
</list-item>
<list-item>
<p>What is the reported efficacy and rate of bacterial degradation across different studies, and what factors (e.g., polymer properties, environmental conditions, bacterial species/consortia) most significantly influence this process?</p>
</list-item>
<list-item>
<p>What are the key challenges and limitations in translating laboratory-based findings into effective, field-scale bioremediation applications?</p>
</list-item>
</list>
</p>
<p>By systematically collating and evaluating the evidence, this review will delineate the current frontiers of knowledge, highlight critical research gaps, and provide an evidence-based foundation to guide future scientific inquiry, policy development, and biotechnological innovation in the fight against microplastic pollution.</p>
</sec>
</sec>
<sec id="s2">
<label>2</label>
<title>Review methodology</title>
<p>This systematic review was conducted in strict accordance with the Preferred Reporting Items for Systematic Reviews and Meta-Analyses (PRISMA 2020) guidelines (<xref ref-type="bibr" rid="B48">Page et al., 2021</xref>) to ensure a comprehensive, transparent, and reproducible synthesis of the global literature on bacterial degradation of microplastics in aquatic environments. The protocol involved a systematic search strategy, a multi-stage screening process guided by pre-defined eligibility criteria, data extraction, risk-of-bias assessment, and qualitative thematic synthesis.</p>
<sec id="s2-1">
<label>2.1</label>
<title>Protocol development and registration</title>
<p>Prior to the commencement of the literature search, a detailed review protocol was developed outlining the objectives, search strategy, inclusion/exclusion criteria, data extraction framework, and synthesis methods. The protocol was designed to minimize selection and publication bias and to ensure reproducibility. While registration in a prospective register such as PROSPERO was considered, the interdisciplinary scope of this review spanning environmental microbiology, biotechnology, and polymer science extended beyond the biomedical focus of such registries. Therefore, an internal protocol was maintained as the guiding document for all subsequent steps.</p>
</sec>
<sec id="s2-2">
<label>2.2</label>
<title>Search strategy and information sources</title>
<p>A systematic literature search was conducted in June 2025 across three major electronic databases: Scopus, Web of Science Core Collection, and PubMed. These databases were selected for their extensive coverage of peer-reviewed literature in environmental science, microbiology, biotechnology, and chemical engineering. The search strategy was constructed around four central conceptual themes: micro-plastics, bacteria, biodegradation, and aquatic environments. A comprehensive set of keywords and Boolean operators (AND, OR) was employed for each concept. The full, detailed search strategy as executed in each database is provided in Supplementary Table S1. For example, the search string used in Scopus was: (micro-plastic OR nano-plastic) AND (bacterial OR microbial) AND (degradation OR biodegradation OR mineral OR de-polymer) AND (aquatic OR marine OR freshwater OR river OR lake).</p>
<p>The search was restricted to peer-reviewed journal articles, systematic reviews, and meta-analyses published in English between 1 January 2000 and 30 June 2025. This date range captures the modern era of micro-plastic research and ensures the review reflects current advancements. Grey literature, preprints, and non-English publications were excluded to maintain consistency in peer-review standards and reproducibility of findings.</p>
</sec>
<sec id="s2-3">
<label>2.3</label>
<title>Study eligibility criteria (PICOS framework)</title>
<p>Study selection was guided by predefined eligibility criteria structured according to the PICOS (Population, Intervention, Comparator, Outcomes, Study design) framework, as detailed in <xref ref-type="table" rid="T1">Table 1</xref>. This framework ensured a focused and relevant selection of studies.</p>
<table-wrap id="T1" position="float">
<label>TABLE 1</label>
<caption>
<p>Study eligibility criteria based on the PICOS framework.</p>
</caption>
<table>
<thead valign="top">
<tr>
<th align="left">PICOS element</th>
<th align="left">Inclusion criteria</th>
<th align="left">Exclusion criteria</th>
</tr>
</thead>
<tbody valign="top">
<tr>
<td align="left">Population</td>
<td align="left">Synthetic polymer microplastics (e.g., PE, PP, PET, PS) in aquatic environments or laboratory simulations thereof</td>
<td align="left">Macroplastics; bioplastics (e.g., PHA, PLA) without synthetic comparison; terrestrial/soil systems</td>
</tr>
<tr>
<td align="left">Intervention</td>
<td align="left">Exposure to or involvement of bacteria (pure cultures, enriched consortia, or natural communities) in degradation</td>
<td align="left">Studies focusing solely on fungal, algal, or abiotic degradation without bacterial component</td>
</tr>
<tr>
<td align="left">Comparator</td>
<td align="left">Compared to untreated/sterile controls, different bacterial strains, or varying environmental conditions</td>
<td align="left">No specific comparator required for inclusion</td>
</tr>
<tr>
<td align="left">Outcomes</td>
<td align="left">Measures of degradation (weight loss, molecular weight reduction, CO<sub>2</sub> production, enzyme activity) or mechanistic characterization</td>
<td align="left">Studies not reporting quantitative or qualitative evidence of degradation or its mechanisms</td>
</tr>
<tr>
<td align="left">Study design</td>
<td align="left">Peer-reviewed journal articles, systematic reviews, meta-analyses</td>
<td align="left">Opinion pieces, editorials, conference abstracts, book chapters, non-peer-reviewed reports</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="s2-4">
<label>2.4</label>
<title>Study selection and screening process</title>
<p>The study selection process followed a rigorous two-stage screening protocol to minimize bias, as illustrated in the PRISMA flow diagram (<xref ref-type="fig" rid="F1">Figure 1</xref>). Initial searches across the databases yielded 639 records. After merging the records, 244 duplicates were identified and removed using a combination of automated tools in EndNote XX and manual verification. This resulted in 395 unique records proceeding to the title and abstract screening stage.</p>
<fig id="F1" position="float">
<label>FIGURE 1</label>
<caption>
<p>PRISMA Flow Diagram illustrating the study selection process.</p>
</caption>
<graphic xlink:href="feart-14-1752600-g001.tif">
<alt-text content-type="machine-generated">Flowchart of systematic review process with four stages: Identification, Screening, Eligibility, and Included. Records were identified from Scopus (294), Web of Science (256), and PubMed (89), totaling 639. After removing 244 duplicates, 395 records were screened. Of these, 241 were excluded. 154 full-text articles were assessed, with 74 excluded for reasons like not meeting PICOS criteria, lack of data, or unavailable full text. Finally, 80 studies were included in the qualitative synthesis.</alt-text>
</graphic>
</fig>
<p>Two independent reviewers screened all 395 records against the eligibility criteria, with any discrepancies resolved through discussion and consensus. During this initial screening, 241 records were excluded as clearly irrelevant to the review&#x2019;s objectives. The full texts of the remaining 154 articles were then retrieved and subjected to a detailed eligibility assessment by the same two reviewers. At this stage, 74 articles were excluded for specific reasons, including failure to meet the PICOS criteria, lack of original data, or unavailability of the full text. Through this stringent process, a total of 80 studies were identified as meeting all inclusion criteria and were therefore selected for data extraction and qualitative synthesis.</p>
</sec>
<sec id="s2-5">
<label>2.5</label>
<title>Data extraction and management</title>
<p>Data from the 80 included studies were systematically extracted using a standardized, pre-piloted form in Microsoft Excel. The extracted information encompassed essential bibliographic details (authors, publication year, journal), key study characteristics (including design, polymer type, bacterial agents, and incubation conditions), and methodological specifics such as the analytical techniques employed.</p>
<p>Additionally, the extraction captured the core research outcomes, including quantitative degradation metrics, identified enzymes or genes, metabolic pathways, and main conclusions, as well as important contextual factors like limitations reported by the authors. All compiled data are presented in Supplementary Table S2, providing a comprehensive overview and enabling effective cross-study comparison. Supplementary Table S2 summarizes key characteristics of the 80 studies included in the systematic review. For each study, information is provided on the polymer type investigated, the bacterial system used (species or consortium), the study design (e.g., laboratory experiment, mesocosm, or field study), the primary degradation metrics reported, and the main findings.</p>
</sec>
<sec id="s2-6">
<label>2.6</label>
<title>Quality assessment and risk of bias in individual studies</title>
<p>The quality and risk of bias of the included primary studies were evaluated using a customized critical appraisal checklist, adapted from established tools for laboratory-based environmental research. This process ensured a systematic and rigorous assessment of the methodological soundness of each study.</p>
<p>Each study was appraised against four key criteria. These were: the adequacy of material characterization for both the polymer and bacterial inoculum; the inclusion of appropriate sterile or abiotic controls; the sufficiency of biological and technical replication; and the analytical rigor demonstrated through the use of multiple, complementary methods to confirm degradation. For each criterion, studies were assigned a rating of low, high, or unclear risk of bias.</p>
<p>The results of this assessment are synthesized visually in <xref ref-type="fig" rid="F2">Figure 2</xref> and detailed in Supplementary Table S3. This evaluation directly informed the critical analysis within the review&#x2019;s discussion (<xref ref-type="sec" rid="s8">Section 8</xref>), serving to highlight the prevailing methodological strengths and limitations across the entire evidence base. <xref ref-type="fig" rid="F2">Figure 2</xref> elucidates the risk of bias assessment summary. The results of this assessment were used to inform the critical analysis of methodological strengths and limitations in <xref ref-type="sec" rid="s8">Section 8</xref>. The below stacked bar chart summarizes the risk of bias across four domains for the 80 included studies. The y-axis shows the percentage of studies in each risk category. The green segments indicate Low Risk, yellow indicate Unclear Risk, and red indicate High Risk (<xref ref-type="fig" rid="F2">Figure 2</xref>).</p>
<fig id="F2" position="float">
<label>FIGURE 2</label>
<caption>
<p>Risk of bias assessment summary.</p>
</caption>
<graphic xlink:href="feart-14-1752600-g002.tif">
<alt-text content-type="machine-generated">Bar chart showing bias domains across four categories: Material Characterization, Use of Controls, Replication, and Analytical Rigor. Each category is divided into Low Risk (green), Unclear Risk (orange), and High Risk (red). Material Characterization has 60% Low, 30% Unclear, 10% High. Use of Controls has 35% Low, 45% Unclear, 20% High. Replication has 20% Low, 40% Unclear, 40% High. Analytical Rigor has 70% Low, 40% Unclear, 10% High.</alt-text>
</graphic>
</fig>
</sec>
<sec id="s2-7">
<label>2.7</label>
<title>Data synthesis and analysis approach (qualitative thematic synthesis)</title>
<p>Due to substantial methodological heterogeneity across the included studies, a quantitative meta-analysis was deemed inappropriate. A critical analysis revealed profound variability across four key domains (<xref ref-type="table" rid="T2">Table 2</xref>), which collectively preclude the statistical pooling of results. These domains are: 1) the polymer substrate&#x2019;s form, crystallinity, and pre-treatment; 2) the biological system, ranging from pure cultures to complex consortia; 3) the incubation conditions, including media, duration, and environmental parameters; and 4) the reported degradation metrics, which employ different analytical endpoints with varying sensitivity. This heterogeneity prevents direct cross-study comparison of quantitative efficacy data. Therefore, a qualitative thematic synthesis approach was employed to integrate complementary evidence, identify consistent trends, and draw comparative insights across these methodologically diverse studies.</p>
<table-wrap id="T2" position="float">
<label>TABLE 2</label>
<caption>
<p>Analysis of methodological heterogeneity across included studies precluding quantitative meta-analysis.</p>
</caption>
<table>
<thead valign="top">
<tr>
<th align="center">Source of heterogeneity</th>
<th align="center">Description of variability</th>
<th align="center">Impact on synthesis</th>
</tr>
</thead>
<tbody valign="top">
<tr>
<td align="center">Polymer Substrate</td>
<td align="left">Form (film, powder, pellet), crystallinity, molecular weight, additive content, and pre-treatment (e.g., UV weathering)</td>
<td align="left">Fundamental differences in substrate accessibility and chemistry make direct comparison of degradation rates invalid</td>
</tr>
<tr>
<td align="center">Biological System</td>
<td align="left">Use of pure cultures vs. enriched consortia vs. natural communities; phylogenetic identity and enzymatic repertoire of degraders</td>
<td align="left">Efficacy is highly strain- and consortium-specific; functional redundancy varies</td>
</tr>
<tr>
<td align="center">Incubation Conditions</td>
<td align="left">Media composition, temperature, pH, duration (days to months), shaking/aeration, and nutrient supplementation</td>
<td align="left">Conditions drastically influence microbial growth and enzymatic activity, leading to non-standardized rate kinetics</td>
</tr>
<tr>
<td align="center">Degradation Metrics</td>
<td align="left">Reliance on different analytical endpoints (e.g., weight loss, SEM, FTIR, GPC, CO<sub>2</sub> evolution) with varying sensitivity and reporting units</td>
<td align="left">Prevents statistical pooling of results; qualitative integration of complementary evidence is required</td>
</tr>
</tbody>
</table>
</table-wrap>
<p>The synthesis of findings was structured according to pre-defined thematic categories, which were directly derived from the research questions and encompassed the diversity and dynamics of plastisphere communities, the molecular and biochemical mechanisms underpinning degradation, the measured efficacy and kinetics of the process under varying conditions, and the persistent challenges in translating laboratory results to field applications.</p>
<p>Findings were tabulated where possible (e.g., <xref ref-type="table" rid="T3">Tables 3</xref>&#x2013;<xref ref-type="table" rid="T9">9</xref>) to facilitate comparison. Subgroup exploration was conducted within homogeneous clusters (e.g., PET degradation by <italic>Ideonella sakaiensis</italic>) to identify trends, but formal statistical pooling was not performed due to inconsistent outcome reporting.</p>
<table-wrap id="T3" position="float">
<label>TABLE 3</label>
<caption>
<p>Synthesis of key bacterial genera implicated in microplastic degradation in aquatic environments.</p>
</caption>
<table>
<thead valign="top">
<tr>
<th align="center">Bacterial genus</th>
<th align="center">Primary polymer targets</th>
<th align="center">Key degradation mechanisms/enzymes</th>
<th align="center">Documented evidence of efficacy</th>
<th align="center">Notes/contextual factors</th>
</tr>
</thead>
<tbody valign="top">
<tr>
<td align="left">
<italic>Pseudomonas</italic>
</td>
<td align="left">Polyethylene (PE), Polystyrene (PS)</td>
<td align="left">Alkane hydroxylases, Laccases, robust biofilm formation</td>
<td align="left">
<italic>P. aeruginosa</italic> DSM 50071: 15% reduction in PE molecular weight in 60 days (<xref ref-type="bibr" rid="B29">Kim et al., 2020</xref>). Frequent dominance in PE/PS plastisphere communities (<xref ref-type="bibr" rid="B19">Howard et al., 2025</xref>)</td>
<td align="left">Metabolically versatile primary colonizer; efficacy is often dependent on polymer oxidation state (e.g., UV-weathered MPs)</td>
</tr>
<tr>
<td align="left">
<italic>Bacillus</italic>
</td>
<td align="left">Polyethylene Terephthalate (PET), Polypropylene (PP)</td>
<td align="left">Cutinase-like hydrolases, Lipases; resilience via endospores</td>
<td align="left">
<italic>B. subtilis</italic> &#x26; <italic>B. cereus</italic>: Hydrolysis of PET ester bonds, releasing terephthalic acid (<xref ref-type="bibr" rid="B58">Roberts et al., 2020</xref>; <xref ref-type="bibr" rid="B28">Khalil et al., 2025</xref>)</td>
<td align="left">Spore formation aids survival in variable environments; promising for bioaugmentation strategies</td>
</tr>
<tr>
<td align="left">Rhodococcus</td>
<td align="left">Polyethylene (PE), Polypropylene (PP)</td>
<td align="left">Alkane metabolism pathways; strong adhesion via mycolic acid in cell wall</td>
<td align="left">
<italic>R. ruber</italic>: Significant biofilm formation and weight loss of PE films (<xref ref-type="bibr" rid="B13">Gorish et al., 2024</xref>)</td>
<td align="left">Particularly effective on hydrophobic polymers due to cell wall structure; specializes in aliphatic chain oxidation</td>
</tr>
<tr>
<td align="left">Ideonella</td>
<td align="left">Polyethylene Terephthalate (PET)</td>
<td align="left">PETase, MHETase (highly specialized enzyme system)</td>
<td align="left">
<italic>I. sakaiensis</italic> 201-F6: Near-complete degradation of low-crystallinity PET films (<xref ref-type="bibr" rid="B77">Yoshida et al., 2016</xref>)</td>
<td align="left">Landmark discovery; represents a highly specialized, but narrow-spectrum, degradation pathway</td>
</tr>
<tr>
<td align="left">Other notable genera (e.g., <italic>Acinetobacter, Comamonas, Streptomyces</italic>)</td>
<td align="left">Various (PE, PP, PS)</td>
<td align="left">Likely a diverse range of oxidative and hydrolytic enzymes; often act in consortia</td>
<td align="left">Recurrent identification in plastisphere metagenomic studies (<xref ref-type="bibr" rid="B67">Szczyrba et al., 2025</xref>)</td>
<td align="left">Their specific roles are often less characterized but are crucial in complex microbial consortia for complete mineralization</td>
</tr>
</tbody>
</table>
</table-wrap>
<table-wrap id="T4" position="float">
<label>TABLE 4</label>
<caption>
<p>Influence of polymer type on plastisphere community structure and degradation pathways.</p>
</caption>
<table>
<thead valign="top">
<tr>
<th align="center">Polymer type</th>
<th align="center">Key chemical/Physical properties</th>
<th align="center">Selected microbial taxa</th>
<th align="center">Dominant degradation mechanism</th>
<th align="center">Degradation kinetics &#x26; challenges</th>
</tr>
</thead>
<tbody valign="top">
<tr>
<td align="left">Polyethylene (PE) &#x26; Polypropylene (PP)</td>
<td align="left">
<list list-type="simple">
<list-item>
<p>&#x2022; C-C backbone (non-hydrolyzable)</p>
</list-item>
<list-item>
<p>&#x2022; High hydrophobicity</p>
</list-item>
<list-item>
<p>&#x2022; High molecular weight</p>
</list-item>
</list>
</td>
<td align="left">
<list list-type="simple">
<list-item>
<p>&#x2022; <italic>Rhodococcus</italic>
</p>
</list-item>
<list-item>
<p>&#x2022; <italic>Pseudomonas</italic>
</p>
</list-item>
<list-item>
<p>&#x2022; Other Gammaproteobacteria (alkane-degraders)</p>
</list-item>
</list>
</td>
<td align="left">Oxidative. Initial oxidation of C-C bonds by enzymes (e.g., hydroxylases) is the rate-limiting step, often preceded/bridged by abiotic weathering (e.g., UV)</td>
<td align="left">Slow. Abiotic weathering often a prerequisite. High molecular weight and crystallinity are major barriers. Complete mineralization is rare in reported studies</td>
</tr>
<tr>
<td align="left">Polyethylene Terephthalate (PET)</td>
<td align="left">
<list list-type="simple">
<list-item>
<p>&#x2022; Ester bonds (hydrolyzable)</p>
</list-item>
<list-item>
<p>&#x2022; Semi-crystalline structure</p>
</list-item>
</list>
</td>
<td align="left">
<list list-type="simple">
<list-item>
<p>&#x2022; <italic>Ideonella</italic>
</p>
</list-item>
<list-item>
<p>&#x2022; <italic>Bacillus</italic>
</p>
</list-item>
<list-item>
<p>&#x2022; <italic>Thermobifida</italic>
</p>
</list-item>
</list>
</td>
<td align="left">Hydrolytic<break/>Direct enzymatic hydrolysis of ester bonds by cutinases, lipases, and PETases</td>
<td align="left">Moderate to fast.<break/>Efficiency highly dependent on crystallinity. Low-crystallinity PET (e.g., from bottles) degrades more readily. Thermophilic enzymes show higher efficacy</td>
</tr>
<tr>
<td align="left">Polystyrene (PS)</td>
<td align="left">
<list list-type="simple">
<list-item>
<p>&#x2022; Aromatic backbone (phenyl rings)</p>
</list-item>
<list-item>
<p>&#x2022; Hydrophobic</p>
</list-item>
</list>
</td>
<td align="left">
<list list-type="simple">
<list-item>
<p>&#x2022; <italic>Pseudomonas</italic>
</p>
</list-item>
<list-item>
<p>&#x2022; <italic>Cupriavidus</italic>
</p>
</list-item>
</list>
</td>
<td align="left">Oxidative (aromatic ring attack)<break/>Dioxygenases target the phenyl ring, leading to ring cleavage and breakdown</td>
<td align="left">Slow. The stable aromatic structure is recalcitrant. Often results in lower microbial diversity but high abundance of specialized degraders. By-products can be toxic <xref ref-type="bibr" rid="B64">Shereen et al. (2025)</xref>
</td>
</tr>
<tr>
<td align="left">General Plastisphere</td>
<td align="center">N/A</td>
<td align="left">Primary Colonizers: <italic>Pseudomonas, Bacillus</italic>
<break/>Secondary Successors: Specialist hydrocarbon-degraders</td>
<td align="left">Consortia-based/successional<break/>Primary colonizers condition the surface and initiate breakdown; secondary successors utilize intermediates, leading to synergistic mineralization</td>
<td align="left">Enhanced in consortia<break/>No single species possesses the full enzymatic repertoire. Biofilm EPS matrix concentrates enzymes and facilitates cross-feeding, improving overall kinetics</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>Degradation kinetics is generalized from laboratory studies and may be slower in complex environmental settings.</p>
</fn>
<fn>
<p>Key: MP, Microplastic; EPS, Extracellular Polymeric Substances; FTIR, Fourier-Transform Infrared Spectroscopy.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<table-wrap id="T5" position="float">
<label>TABLE 5</label>
<caption>
<p>Key enzymatic systems in bacterial degradation of microplastics.</p>
</caption>
<table>
<thead valign="top">
<tr>
<th align="center">Enzyme class</th>
<th align="center">Representative enzymes</th>
<th align="center">Primary polymer targets</th>
<th align="center">Catalytic action</th>
<th align="center">Key breakdown products</th>
<th align="center">Notable bacterial sources</th>
</tr>
</thead>
<tbody valign="top">
<tr>
<td align="left">Hydrolases</td>
<td align="left">PETase, cutinase, lipase, esterase</td>
<td align="left">PET, PLA, PCL</td>
<td align="left">Hydrolysis of ester bonds</td>
<td align="left">MHET, terephthalic acid, ethylene glycol, oligomers</td>
<td align="left">Ideonella sakaiensis, Thermobifida spp., <italic>Bacillus</italic> spp.</td>
</tr>
<tr>
<td align="left">Oxidoreductases</td>
<td align="left">Alkane hydroxylase, monooxygenase</td>
<td align="left">PE, PP</td>
<td align="left">Oxidation of C-C and C-H bonds, introducing carbonyl/hydroxyl groups</td>
<td align="left">Alcohols, aldehydes, carboxylic acids</td>
<td align="left">
<italic>Pseudomonas</italic> spp., Rhodococcus spp.</td>
</tr>
<tr>
<td align="left">Oxidoreductases</td>
<td align="left">Dioxygenase</td>
<td align="left">PS (Aromatic ring)</td>
<td align="left">Cleavage of the aromatic benzene ring</td>
<td align="left">Styrene oxide, phenylacetaldehyde, pyruvate</td>
<td align="left">
<italic>Pseudomonas</italic> spp., Cupriavidus spp.</td>
</tr>
<tr>
<td align="left">Oxidoreductases</td>
<td align="left">Laccase (with/without mediators)</td>
<td align="left">PE, PS, PVC</td>
<td align="left">Radical formation leading to chain depolymerization</td>
<td align="left">Various radical intermediates, fragmented oligomers</td>
<td align="left">
<italic>Bacillus</italic> spp., <italic>Streptomyces</italic> spp.</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>Key: PE, Polyethylene; PP, Polypropylene; PET, Polyethylene Terephthalate; PS, Polystyrene; PVC, Polyvinyl Chloride; PLA, Polylactic Acid; PCL, Polycaprolactone; MHET, mono(2-hydroxyethyl) terephthalic acid.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<table-wrap id="T6" position="float">
<label>TABLE 6</label>
<caption>
<p>Synthesis of degradation efficacy for common plastic polymers by key bacterial species/consortia.</p>
</caption>
<table>
<thead valign="top">
<tr>
<th align="center">Polymer type</th>
<th align="center">Representative bacterial system</th>
<th align="center">Incubation duration</th>
<th align="center">Key efficacy metrics &#x26; results</th>
<th align="center">Influencing factors &#x26; notes</th>
</tr>
</thead>
<tbody valign="top">
<tr>
<td align="left">Polyethylene (PE)</td>
<td align="left">Rhodococcus ruber (strain C208)</td>
<td align="center">60 days</td>
<td align="left">&#x223c;8% weight loss; Significant biofilm formation; Increase in carbonyl index (FTIR)</td>
<td align="left">Pre-oxidation (e.g., UV weathering) is a critical prerequisite for significant degradation</td>
</tr>
<tr>
<td align="left">Polyethylene (PE)</td>
<td align="left">Consortium (<italic>Pseudomonas</italic> spp. &#x2b; <italic>Bacillus</italic> spp.)</td>
<td align="center">120 days</td>
<td align="left">&#x223c;12% weight loss; Reduction in molecular weight (Mw) by 20% (GPC) <xref ref-type="bibr" rid="B53">Prata et al. (2024)</xref>
</td>
<td align="left">Consortia show synergistic effects, leading to higher efficacy than individual strains</td>
</tr>
<tr>
<td align="left">Polypropylene (PP)</td>
<td align="left">
<italic>Bacillus</italic> cereus</td>
<td align="center">90 days</td>
<td align="left">&#x223c;4% weight loss; Surface pitting and erosion (SEM)</td>
<td align="left">Generally slower than PE degradation due to methyl groups increasing recalcitrance</td>
</tr>
<tr>
<td align="left">Polyethylene Terephthalate (PET)</td>
<td align="left">Ideonella sakaiensis 201-F6</td>
<td align="center">42 days</td>
<td align="left">Near-complete degradation of low-crystallinity PET film; Clear zone formation on agar <xref ref-type="bibr" rid="B32">Kumar et al. (2022)</xref>
</td>
<td align="left">Highly effective but specific to low-crystallinity PET (e.g., from bottles)</td>
</tr>
<tr>
<td align="left">Polyethylene Terephthalate (PET)</td>
<td align="left">Thermobifida fusca cutinase</td>
<td align="center">96 h (at 55 &#xb0;C)</td>
<td align="left">50% mass loss of amorphous PET; Release of TPA and EG (HPLC)</td>
<td align="left">Thermophilic enzymes show superior performance due to increased polymer mobility</td>
</tr>
<tr>
<td align="left">Polystyrene (PS)</td>
<td align="left">
<italic>Pseudomonas aeruginosa</italic>
</td>
<td align="center">60 days</td>
<td align="left">&#x223c;2% weight loss; Breakdown of aromatic rings detected (FTIR); CO<sub>2</sub> evolution confirmed</td>
<td align="left">Degradation is slow; often limited to the formation of oxidized oligomers</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>Abbreviations: FTIR, Fourier-Transform Infrared Spectroscopy; GPC, Gel Permeation Chromatography; SEM, Scanning Electron Microscopy; HPLC, High-Performance Liquid Chromatography; TPA, Terephthalic Acid; EG, Ethylene Glycol.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<table-wrap id="T7" position="float">
<label>TABLE 7</label>
<caption>
<p>Critical barriers to field application of bacterial microplastic degradation.</p>
</caption>
<table>
<thead valign="top">
<tr>
<th align="center">Barrier category</th>
<th align="center">Specific challenge</th>
<th align="center">Impact on bioremediation efficacy</th>
<th align="center">Current research gaps</th>
</tr>
</thead>
<tbody valign="top">
<tr>
<td align="left">Ecological</td>
<td align="left">Competition with indigenous microbiota</td>
<td align="left">Reduces survival and activity of introduced degraders (bio-augmentation)</td>
<td align="left">Strategies to enhance ecological competence of degraders; understanding keystone species in plastisphere</td>
</tr>
<tr>
<td align="left">Ecological</td>
<td align="left">Predation (e.g., by protozoa)</td>
<td align="left">Rapid removal of bacterial inoculants from the water column</td>
<td align="left">Development of predation-resistant strains or delivery mechanisms (e.g., microencapsulation)</td>
</tr>
<tr>
<td align="left">Environmental</td>
<td align="left">Catabolite repression</td>
<td align="left">Preferential consumption of natural organic matter over microplastics</td>
<td align="left">Understanding regulatory networks in complex nutrient environments</td>
</tr>
<tr>
<td align="left">Engineering &#x26; economic</td>
<td align="left">Scalability of inoculant production &#x26; delivery</td>
<td align="left">High cost and logistical infeasibility for large-scale application</td>
<td align="left">Development of low-cost, high-yield cultivation and stable, slow-release formulation technologies</td>
</tr>
<tr>
<td align="left">Methodological</td>
<td align="left">Monitoring and verification in open systems</td>
<td align="left">Inability to accurately attribute microplastic removal to the biological intervention</td>
<td align="left">Development of isotopically labelled (&#x201c;C-13&#x201d;) plastics and standardized molecular tools for tracking specific degradative activity</td>
</tr>
</tbody>
</table>
</table-wrap>
<table-wrap id="T8" position="float">
<label>TABLE 8</label>
<caption>
<p>Comparative analysis of application pathways for plastic-degrading bacteria.</p>
</caption>
<table>
<thead valign="top">
<tr>
<th align="center">Application pathway</th>
<th align="center">Technological readiness</th>
<th align="center">Key advantage</th>
<th align="center">Primary challenge</th>
<th align="center">Most viable polymer targets</th>
</tr>
</thead>
<tbody valign="top">
<tr>
<td align="left">
<italic>In-situ</italic> bioremediation (environmental cleanup)</td>
<td align="center">Low (conceptual/experimental)</td>
<td align="left">Targets dispersed pollution in the environment</td>
<td align="left">Formidable ecological, monitoring, and scalability barriers; regulatory hurdles</td>
<td align="left">Weathered PE/PP (slow); PS (experimental)</td>
</tr>
<tr>
<td align="left">
<italic>Ex-situ</italic> bioremediation (contained systems)</td>
<td align="left">Medium (pilot-scale)</td>
<td align="left">Controlled conditions enhance efficacy and monitoring</td>
<td align="left">Requires collection and concentration of waste; energy input for operation</td>
<td align="left">PET microfibers in wastewater; mixed plastic waste in bioreactor landfills</td>
</tr>
<tr>
<td align="left">Industrial bioprocessing (biorecycling)</td>
<td align="left">High (pilot to early commercial)</td>
<td align="left">Enables circular economy; produces valuable monomers; economically incentivized</td>
<td align="left">High capital cost; competition with virgin plastic production; requires efficient waste sorting</td>
<td align="left">PET, PLA, and other hydrolysable polymers</td>
</tr>
</tbody>
</table>
</table-wrap>
<table-wrap id="T9" position="float">
<label>TABLE 9</label>
<caption>
<p>Critical knowledge gaps and recommended research directions.</p>
</caption>
<table>
<thead valign="top">
<tr>
<th align="center">Knowledge gap category</th>
<th align="center">Specific gap</th>
<th align="center">Implications</th>
<th align="center">Recommended research direction</th>
</tr>
</thead>
<tbody valign="top">
<tr>
<td align="left">Environmental efficacy &#x26; fate</td>
<td align="left">Long-term fate and efficacy in complex matrices (sediment, water column)</td>
<td align="left">Unknown if degradation continues or stalls; inability to model environmental impact</td>
<td align="left">Long-term mesocosm studies with environmentally-weathered MPs and native microbial communities</td>
</tr>
<tr>
<td align="left">Ecological impact</td>
<td align="left">Ecotoxicity of plastic additives and degradation by-products</td>
<td align="left">Bioremediation may solve one problem (MPs) but create another (toxic leachate)</td>
<td align="left">Integrated chemical and toxicological assessments (e.g., using bioassays) alongside degradation studies</td>
</tr>
<tr>
<td align="left">Technology development</td>
<td align="left">Design and ecology of engineered consortia</td>
<td align="left">Single strains are ineffective; random consortia are unpredictable</td>
<td align="left">Use of synthetic ecology principles to design stable, cooperative, and effective multi-species consortia</td>
</tr>
<tr>
<td align="left">Monitoring &#x26; verification</td>
<td align="left">Tools for <italic>in-situ</italic> tracking of degradation activity</td>
<td align="left">Inability to validate bioremediation efforts in the field</td>
<td align="left">Development of molecular probes (e.g., for degradative gene expression) and isotopic tracer (<sup>13</sup>C-MP) methods</td>
</tr>
<tr>
<td align="left">Polymer-microbe specificity</td>
<td align="left">Systematic understanding of how polymer properties govern microbial colonization and degradation</td>
<td align="left">Inefficient &#x201c;trial-and-error&#x201d; approach to finding degraders</td>
<td align="left">High-throughput screening coupled with detailed polymer analytics to build predictive models</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
</sec>
<sec id="s3">
<label>3</label>
<title>The plastisphere: diversity, dynamics, and polymer-specific selection</title>
<p>The &#x201c;plastisphere,&#x201d; a unique microbial habitat that forms on plastic debris, serves as a critical interface where biodegradation is initiated and modulated (<xref ref-type="bibr" rid="B7">Bocci et al., 2024</xref>). Our synthesis reveals that its assembly is a non-random, sequential process governed by polymer chemistry, which acts as a primary environmental filter selecting for distinct microbial consortia (<xref ref-type="bibr" rid="B4">Battulga et al., 2024</xref>).</p>
<sec id="s3-1">
<label>3.1</label>
<title>Key bacterial taxa and their degradative niches</title>
<p>Analysis confirms that specific bacterial genera are consistently enriched on microplastics (MPs) across diverse aquatic habitats due to their metabolic versatility and adaptive traits (<xref ref-type="bibr" rid="B63">Scott et al., 2025</xref>). Notable primary colonizers include <italic>Pseudomonas</italic> spp., which dominate on polyethylene and polystyrene via robust biofilm formation and the secretion of nonspecific oxidoreductases (<xref ref-type="bibr" rid="B78">Zhang et al., 2022</xref>). Similarly, <italic>Bacillus</italic> spp., resilient through endospore formation, is frequently associated with polyethylene terephthalate and polypropylene through their secretion of hydrolytic enzymes (<xref ref-type="bibr" rid="B16">Gupta and Devi, 2020</xref>). <italic>Rhodococcus</italic> spp. also shows a strong affinity for polyolefins like polyethylene and polypropylene, leveraging hydrophobic cell walls and established alkane metabolic pathways for polymer oxidation (<xref ref-type="bibr" rid="B74">Yadav et al., 2025</xref>).</p>
<p>Beyond these well-known genera, specialized and emerging taxa highlight advanced degradative strategies. The landmark discovery of <italic>Ideonella sakaiensis</italic> 201-F6, which utilizes the specialized PETase and MHETase enzyme system, demonstrates highly efficient, polymer-specific hydrolysis (<xref ref-type="bibr" rid="B76">Yip et al., 2024</xref>). Furthermore, recurrent identification of genera such as <italic>Acinetobacter</italic>, <italic>Comamonas</italic>, and <italic>Streptomyces</italic> in plastisphere studies indicates their significant, though often less characterized, roles within complex microbial consortia dedicated to degradation (<xref ref-type="bibr" rid="B7">Bocci et al., 2024</xref>).</p>
</sec>
<sec id="s3-2">
<label>3.2</label>
<title>Integrated dynamics: biofilm succession and polymer-driven selection</title>
<p>Colonization begins with pioneering bacteria (e.g., <italic>Pseudomonas</italic>, <italic>Bacillus</italic>) adhering to the conditioned &#x201c;eco-corona&#x201d; of the MP. They secrete extracellular polymeric substances (EPS), forming a biofilm matrix that concentrates enzymes and facilitates microbial cross-feeding. As degradation initiates, the microenvironment changes; driving microbial succession (<xref ref-type="bibr" rid="B62">Scheidweiler et al., 2019</xref>). Secondary colonizers, often specialists in utilizing intermediary breakdown products, establish themselves, leading to a more complex and functionally synergistic consortium crucial for complete mineralization (<xref ref-type="bibr" rid="B4">Battulga et al., 2024</xref>; <xref ref-type="bibr" rid="B7">Bocci et al., 2024</xref>).</p>
<p>Critically, the chemical composition of the polymer acts as a strong selective pressure, directly shaping the microbial community dynamics on its surface (<xref ref-type="bibr" rid="B4">Battulga et al., 2024</xref>). Non-hydrolyzable, hydrophobic polymers like polyethylene (PE) and polypropylene (PP) select for communities dominated by <italic>Rhodococcus</italic> and <italic>Pseudomonas</italic>, which possess oxidative enzymatic systems for the slow oxidation of C-C bonds (<xref ref-type="bibr" rid="B52">Porter et al., 2023</xref>). In contrast, the presence of hydrolysable ester bonds in polyethylene terephthalate (PET) enriches for a distinct community with bacteria like <italic>Ideonella sakaiensis</italic>, <italic>Bacillus</italic>, and <italic>Thermobifida</italic> species that secrete potent hydrolases, particularly when polymer crystallinity is reduced. Meanwhile, the aromatic backbone of polystyrene (PS) presents a unique challenge, often resulting in a lower-diversity community dominated by specific <italic>Pseudomonas</italic> and <italic>Cupriavidus</italic> species capable of aromatic ring cleavage via dioxygenases. This polymer-driven niche differentiation underscores that the plastisphere is not a single entity but a collection of polymer-specific habitats (<xref ref-type="bibr" rid="B44">Mulky et al., 2025</xref>). Consequently, successful bioremediation strategies must be &#x201c;tailored,&#x201d; matching specific bacterial consortia or enzymes to the predominant polymer pollutant. This relationship between polymer type, microbial community structure, and the resulting degradation pathway is illustrated in the accompanying tables (<xref ref-type="table" rid="T3">Tables 3</xref>, <xref ref-type="table" rid="T4">4</xref>).</p>
</sec>
</sec>
<sec id="s4">
<label>4</label>
<title>Molecular and biochemical mechanisms of bacterial degradation</title>
<sec id="s4-1">
<label>4.1</label>
<title>Enzymatic machinery: a systematic overview of hydrolases, oxidoreductases, and laccases</title>
<p>The systematic analysis of the included literature reveals that bacterial degradation of microplastics is primarily mediated by a limited yet versatile set of extracellular enzymes, which can be broadly categorized based on their catalytic mechanism and polymer target, and this class of enzymes is paramount for the degradation of polymers containing hydrolysable bonds, such as polyesters, where the discovery of PETase from <italic>Ideonella sakaiensis</italic> represents a landmark finding, as this enzyme specifically targets the aromatic ester bonds in polyethylene terephthalate (PET), yielding mono(2-hydroxyethyl) terephthalic acid (MHET) as a primary product (<xref ref-type="bibr" rid="B27">Khairul Anuar et al., 2022</xref>; <xref ref-type="bibr" rid="B80">Zhang, X. et al., 2024</xref>), and subsequently, MHETase further hydrolyzes MHET into the monomers terephthalic acid and ethylene glycol (<xref ref-type="bibr" rid="B50">Palit et al., 2025</xref>), while beyond this specialized system, more ubiquitous hydrolytic enzymes like cutinases (from <italic>Thermobifida</italic> and <italic>Bacillus</italic> spp.) and lipases have demonstrated significant efficacy against PET and other aliphatic polyesters by attacking their ester linkages (<xref ref-type="bibr" rid="B40">Maurya et al., 2020</xref>), with the effectiveness of hydrolases being highly dependent on polymer accessibility, as amorphous regions are degraded preferentially over crystalline ones.</p>
<p>For non-hydrolyzable polymers like polyethylene (PE) and polypropylene (PP), which possess inert C-C backbones, oxidative cleavage is the initial and rate-limiting step, where enzymes such as alkane hydroxylases (e.g., from <italic>Pseudomonas</italic> and <italic>Rhodococcus</italic>) introduce hydroxyl groups into the polymer chain, making it more susceptible to further oxidation (<xref ref-type="bibr" rid="B30">Kim et al., 2025</xref>), and monooxygenases and dioxygenases play a critical role, particularly in attacking the aromatic rings of polystyrene (PS), as demonstrated by the styrene degradation pathway involving styrene monooxygenase (SMO) and styrene oxide isomerase, which has been implicated in the breakdown of PS oligomers (<xref ref-type="bibr" rid="B47">Oelschl&#xe4;gel et al., 2018</xref>).</p>
<p>These multi-copper oxidases are noted for their broad substrate specificity and ability to act on a range of polymers, including PE, PS, and polyvinyl chloride (PVC), where laccases catalyze the one-electron oxidation of various substrates, often generating reactive radicals that can lead to polymer chain scission, and their activity is often enhanced in the presence of redox mediators, which can expand their target range (<xref ref-type="bibr" rid="B36">Liu X-h et al., 2025</xref>), and while more commonly associated with fungi, bacterial laccases (e.g., from <italic>Bacillus</italic> spp.) are increasingly being recognized for their role in the plastisphere, with <xref ref-type="table" rid="T5">Table 5</xref> providing a systematic summary of these key enzymatic systems, their targets, and the resulting breakdown products.</p>
</sec>
<sec id="s4-2">
<label>4.2</label>
<title>Metabolic pathways and genetic regulation of polymer breakdown</title>
<p>The initial enzymatic attack generates intermediary compounds that are subsequently assimilated into central metabolic pathways, and our synthesis indicates that bacteria commonly achieve this by co-opting pre-existing catabolic routes originally evolved for natural hydrocarbons and aromatic compounds, as, for example, the degradation of polyethylene terephthalate (PET) yields the monomers terephthalic acid and ethylene glycol, which are then processed via dedicated bacterial pathways for terephthalic acid degradation and glycolytic metabolism, respectively, ultimately feeding into the tricarboxylic acid (TCA) cycle, and in the case of polyolefins like polyethylene (PE) and polypropylene (PP), oxidized products such as fatty alcohols and acids are typically channeled into the &#x3b2;-oxidation cycle to generate acetyl-CoA, while polystyrene (PS) degradation is more complex, with evidence pointing to ring cleavage products entering central metabolism through modified routes of styrene catabolism, and a critical finding is the importance of consortia-based metabolism and cross-feeding, where no single bacterium may possess the complete enzymatic suite for full mineralization (<xref ref-type="bibr" rid="B61">Salinas et al., 2024</xref>; <xref ref-type="bibr" rid="B17">Hakkarainen and Albertsson, 2026</xref>), meaning the breakdown products from one species serve as substrates for another, leading to synergistic degradation more efficient than any single isolate.</p>
<p>This degradative process is an active, regulated cellular response, as genomic and transcriptomic studies confirm that genes encoding key enzymes (e.g., petase, mhetase, alkane hydroxylase operons) are often located on mobile genetic elements, suggesting potential for horizontal gene transfer (<xref ref-type="bibr" rid="B75">Yang et al., 2023</xref>; <xref ref-type="bibr" rid="B76">Yip et al., 2024</xref>), and their expression is tightly regulated and induced by the presence of plastic polymers or their breakdown intermediates, with the molecular response to plastic exposure being an active, regulated process, and genomic and transcriptomic studies included in this review consistently show that the genes encoding the key degradative enzymes are often located on plasmids or genomic islands, suggesting a potential for horizontal gene transfer (<xref ref-type="bibr" rid="B25">Julius et al., 2025</xref>).</p>
<p>The expression of these genes is tightly regulated, as, for instance, the expression of petase and mhetase in <italic>I. sakaiensis</italic> is induced in the presence of PET or its intermediate MHET, and similarly, the genes for alkane hydroxylation in <italic>Pseudomonas</italic> spp. are often part of operons (e.g., the alk system) whose expression is upregulated when the bacterium encounters long-chain alkanes, a regulatory logic it apparently applies to polyethylene (<xref ref-type="bibr" rid="B73">Xiang et al., 2023</xref>), which confirms that bacterial degradation is not a passive process but an active metabolic response to the availability of plastic as a potential carbon source, and <xref ref-type="fig" rid="F3">Figure 3</xref> provides a conceptual summary of the key degradation pathways for the most common plastic polymers, integrating the enzymatic, metabolic, and genetic concepts discussed.</p>
<fig id="F3" position="float">
<label>FIGURE 3</label>
<caption>
<p>Integrated molecular mechanisms of bacterial degradation for major plastic polymers.</p>
</caption>
<graphic xlink:href="feart-14-1752600-g003.tif">
<alt-text content-type="machine-generated">Flowchart depicting the metabolic pathways of polyethylene (PE), polyethylene terephthalate (PET), and polystyrene (PS). Each polymer undergoes enzymatic conversion: PE via oxidoreductases to oxygenated intermediates; PET through hydrolases to monomers and oligomers; PS by dioxygenases to cleaved intermediates. All pathways lead to central metabolism (TCA cycle) and gene regulation, resulting in CO2, H2O, and biomass.</alt-text>
</graphic>
</fig>
</sec>
</sec>
<sec id="s5">
<label>5</label>
<title>Efficacy and kinetics of degradation: evidence from laboratory studies</title>
<p>While the molecular mechanisms provide a blueprint for biodegradation, the practical feasibility of this process hinges on its <italic>efficacy</italic> and <italic>kinetics</italic> under controlled and environmentally relevant conditions. This section synthesizes quantitative evidence from laboratory studies to critically evaluate the rates of degradation, the key factors influencing these rates, and the comparative performance across different bacterial-polymer combinations. Acknowledging the methodological heterogeneity across studies, this synthesis focuses on identifying consistent trends and establishing a realistic benchmark for the current state of the technology.</p>
<sec id="s5-1">
<label>5.1</label>
<title>Quantifying degradation: metrics for weight loss, surface erosion, and CO<sub>2</sub> production</title>
<p>The included studies employ a suite of complementary analytical techniques to quantify biodegradation, as no single metric provides a complete assessment. Direct gravimetric analysis of weight loss remains a fundamental, widely reported measure, though it is often slow to manifest significant change, particularly for recalcitrant polymers like polyolefins. Consequently, more sensitive surface characterization methods are crucial for detecting early-stage degradation (<xref ref-type="bibr" rid="B80">Zhang Z. et al., 2024</xref>). Scanning Electron Microscopy (SEM) provides visual evidence of physical erosion, while Fourier-Transform Infrared Spectroscopy (FTIR) identifies key chemical changes, such as carbonyl formation on polyethylene or the reduction of ester bonds in PET (<xref ref-type="bibr" rid="B3">Azeez and Shenbagaraman, 2025</xref>).</p>
<p>The most definitive proof of complete biodegradation is the measurement of polymer carbon mineralization into CO<sub>2</sub> via respirometric assays, which serves as a gold-standard metric despite often revealing very low conversion rates on laboratory timescales (<xref ref-type="bibr" rid="B15">Guo et al., 2010</xref>). Complementing these approaches, Gel Permeation Chromatography (GPC) offers critical insight into the depolymerization process by detecting reductions in average molecular weight, a clear indicator of chain scission that can occur prior to measurable mass loss. Our synthesis confirms that the most compelling studies utilize this multi-method approach to build a robust, convergent body of evidence (<xref ref-type="bibr" rid="B60">Rydholm et al., 2006</xref>).</p>
</sec>
<sec id="s5-2">
<label>5.2</label>
<title>The impact of environmental factors: temperature, pH, and nutrient availability</title>
<p>The efficacy of bacterial degradation is not an intrinsic property but is profoundly modulated by environmental conditions, with the synthesized evidence revealing several key determinants, where temperature exerts a dual effect, influencing both microbial metabolic rates and polymer physical properties, as, for instance, the activity of PET-degrading enzymes from <italic>Ideonella sakaiensis</italic> and <italic>Thermobifida</italic> spp. is significantly higher at mesophilic (25 &#xb0;C&#x2013;37 &#xb0;C) and thermophilic (55 &#xb0;C&#x2013;70 &#xb0;C) ranges, respectively, and higher temperatures also increase polymer chain mobility, particularly above the glass transition temperature (Tg), making the material more accessible to enzymatic attack (<xref ref-type="bibr" rid="B1">Akram et al., 2024</xref>).</p>
<p>Furthermore, enzyme activity is pH-dependent, with most reported bacterial hydrolases and oxidoreductases operating optimally in a neutral to slightly alkaline pH range (7.0&#x2013;9.0), and significant deviations from this range can inhibit microbial growth and enzymatic efficiency, limiting degradation in highly acidic or alkaline environments (<xref ref-type="bibr" rid="B6">B&#x142;o&#x144;ska et al., 2015</xref>); additionally, nutrient availability has complex, context-dependent effects, where while essential for microbial growth, easily metabolizable co-substrates can lead to catabolite repression, causing bacteria to preferentially consume simple nutrients over the complex polymer, but conversely, in nutrient-poor oligotrophic environments (which mimic many aquatic systems), the plastic may serve as a crucial carbon source, potentially enhancing its biodegradation (<xref ref-type="bibr" rid="B21">Jeske and Gallert, 2021</xref>).</p>
<p>A central finding of this systematic review is the vast disparity in degradation efficacy and kinetics across different polymer-bacteria systems, a central finding of this systematic review is the vast disparity in degradation efficacy and kinetics across different polymer-bacteria systems, and this variability is quantitatively summarized and graphically compared in the accompanying <xref ref-type="table" rid="T6">Table 6</xref> and <xref ref-type="fig" rid="F4">Figure 4</xref>.</p>
<fig id="F4" position="float">
<label>FIGURE 4</label>
<caption>
<p>Comparative kinetics and key determinants of bacterial degradation for major polymer types. <bold>(A)</bold> Comparative dergration kinetics. <bold>(B)</bold> Impact of environmental factors. <bold>(C)</bold> Consorsium vs. single isolate eficiacy. <bold>(D)</bold> Single, isolate and microbial consortium.</p>
</caption>
<graphic xlink:href="feart-14-1752600-g004.tif">
<alt-text content-type="machine-generated">Panel A shows a line graph of cumulative plastic degradation over three hundred days comparing PET of low and high crystallinity and PE/PP/PS, with low crystallinity PET degrading fastest. Panel B is a grid illustrating microbial degradation efficacy under different nutrient and temperature conditions, noting highest efficacy in nutrient-rich, high-temperature environments. Panel C summarizes low degradation efficacy in the open ocean. Panel D is a bar graph showing significantly higher PE degradation by a microbial consortium than a single microbial isolate over ninety days.</alt-text>
</graphic>
</fig>
</sec>
</sec>
<sec id="s6">
<label>6</label>
<title>From lab to field: challenges in environmental application</title>
<p>The promising results from controlled laboratory studies, as synthesized in previous sections, necessitate a critical evaluation of their translation to complex, open environments.</p>
<sec id="s6-1">
<label>6.1</label>
<title>The bio-augmentation vs. bio-stimulation debate: evidence from mesocosm studies</title>
<p>A primary strategic question in bioremediation is whether to introduce specialized degraders (bio-augmentation) or to enhance the activity of the indigenous microbial community (bio-stimulation). Evidence from mesocosm studies which bridge the gap between lab flasks and natural ecosystems provides critical, albeit mixed, insights.</p>
<sec id="s6-1-1">
<label>6.1.1</label>
<title>Bio-augmentation</title>
<p>This approach involves introducing non-native, high-performing bacterial strains (e.g., <italic>Ideonella sakaiensis</italic> for PET or engineered consortia). Mesocosm studies reveal that while bio-augmentation can initially accelerate degradation, the introduced strains often fail to compete with the established, adapted indigenous microbiota, leading to a rapid decline in their population and efficacy (<xref ref-type="bibr" rid="B38">Marquiegui Alvaro et al., 2025</xref>). Factors such as predation, nutrient competition, and unfavorable environmental conditions significantly limit their long-term persistence and activity.</p>
</sec>
<sec id="s6-1-2">
<label>6.1.2</label>
<title>Bio-stimulation</title>
<p>This strategy focuses on manipulating the environment to favor native plastic-degrading microbes by adding rate-limiting nutrients (e.g., nitrogen, phosphorus), oxygen, or other electron acceptors. Evidence suggests this can be a more sustainable approach, as it leverages the existing, adapted microbial network. For instance, the addition of specific nitrogen sources in sediment mesocosms was shown to enhance the natural degradation of weathered PE by stimulating a native consortium of <italic>Pseudomonas</italic> and <italic>Rhodococcus</italic> (<xref ref-type="bibr" rid="B43">Mohamed and Samer, 2023</xref>). The synthesized evidence indicates that a hybrid strategy may be most effective: a one-time, low-dose bio-augmentation with robust, ecologically competent strains, combined with targeted bio-stimulation to support both the introduced and indigenous degraders (<xref ref-type="bibr" rid="B45">Nayak et al., 2021</xref>).</p>
</sec>
</sec>
<sec id="s6-2">
<label>6.2</label>
<title>Critical barriers: competition with indigenous microbes, scalability, and monitoring</title>
<p>The transition from successful mesocosm experiments to field-scale application is hindered by several critical, interconnected barriers, including ecological competition and predation, where in a natural environment, introduced or stimulated degraders must contend with a highly diverse and competitive microbial community, facing pressure from bacteriophages and protozoan grazing, which can rapidly decimate their populations, and where the presence of more easily degradable organic carbon can lead to catabolite repression, causing microbes to preferentially metabolize simple substrates over complex polymers (<xref ref-type="bibr" rid="B32">Kumar et al., 2022</xref>).</p>
<p>Scalability and cost present further monumental challenges, as laboratory processes are optimized in small, homogeneous volumes, and scaling these processes to treat vast, heterogeneous aquatic environments like oceans or lakes presents monumental engineering and economic challenges, making the production, stabilization, and uniform dispersal of effective bacterial consortia or nutrients across square kilometers currently impractical and prohibitively expensive (<xref ref-type="bibr" rid="B9">Crater and Lievense, 2018</xref>).</p>
<p>Finally, monitoring and verification are exceptionally difficult, as accurately measuring the efficacy of a bioremediation intervention in a dynamic open environment requires distinguishing the microplastic mass loss due to the specific intervention from background processes like fragmentation, sedimentation, and transport, which is a major methodological hurdle, highlighting the pressing need for the development of robust tracer techniques and standardized monitoring protocols to validate <italic>in-situ</italic> degradation (<xref ref-type="bibr" rid="B70">Vukovi&#x107; Domanovac, et al., 2025</xref>), with <xref ref-type="table" rid="T7">Table 7</xref> summarizing these key challenges and the associated research gaps that need to be addressed.</p>
<p>The transition from controlled experiments to field-scale application is hindered by profound ecological and practical barriers. A major, often understated, challenge is the paradox of nutrient availability. In oligotrophic open waters, MPs may serve as a crucial carbon source, potentially enhancing biodegradation. However, in nutrient-rich coastal or estuarine environments, the presence of easily metabolizable organic matter can lead to catabolite repression, where microbes preferentially consume simple substrates over complex polymers, effectively halting targeted degradation efforts (<xref ref-type="bibr" rid="B42">Mishra et al., 2025</xref>). Withal, the ecological competence of introduced strains is frequently overestimated. Native microbial communities possess established interactions and defense mechanisms; introduced degraders must compete for space and resources while facing pressure from protozoan grazing and bacteriophage predation, which can rapidly, decimate their populations. This enliven that efficacy in a flask is a poor predictor of environmental persistence and impact.</p>
</sec>
<sec id="s6-3">
<label>6.3</label>
<title>Case studies of promising <italic>in-situ</italic> and <italic>ex-situ</italic> bioremediation attempts</title>
<p>Despite the challenges, a limited number of pioneering studies have attempted to demonstrate bioremediation in real-world contexts, providing valuable proof-of-concept insights.</p>
<sec id="s6-3-1">
<label>6.3.1</label>
<title>
<italic>In-situ</italic> attempts (treatment in the environment)</title>
<p>A notable field trial in a contaminated marina involved the deployment of buoys designed to slowly release a nutrient mix (N, P, K) to stimulate the native microbiome around plastic debris. After 6 months, microbiological analysis showed a significant enrichment of known hydrocarbon-degrading bacteria on the plastic surfaces in the treated area compared to a control. However, direct quantification of MP mass loss was inconclusive, underscoring the monitoring challenge (<xref ref-type="bibr" rid="B53">Prata et al., 2024</xref>).</p>
</sec>
<sec id="s6-3-2">
<label>6.3.2</label>
<title>
<italic>Ex-situ</italic> attempts (treatment in controlled, contained systems)</title>
<p>More success has been reported with <italic>ex-situ</italic> bioreactors, which exemplify a key stage in the translational workflow. A study treating wastewater effluent high in PET microfibers used a dedicated bioreactor inoculated with a defined consortium of <italic>Bacillus</italic> and <italic>Thermobifida</italic> species. Under controlled temperature and aeration, the system achieved a 75% reduction in PET microfiber concentration over 30 days, confirmed by FTIR and SEM analysis (<xref ref-type="bibr" rid="B12">Dhaka et al., 2025</xref>). This highlights the greater controllability and higher efficacy of <italic>ex-situ</italic> systems, making them a more near-term solution for point sources like wastewater treatment plants, and represents a critical step in the pathway from laboratory discovery to field application as conceptualized in <xref ref-type="fig" rid="F5">Figure 5</xref>.</p>
<fig id="F5" position="float">
<label>FIGURE 5</label>
<caption>
<p>Conceptual workflow for transitioning bacterial degradation from laboratory to field application.</p>
</caption>
<graphic xlink:href="feart-14-1752600-g005.tif">
<alt-text content-type="machine-generated">Bioremediation Strategy Development Pipeline diagram outlining three stages: Laboratory Discovery, Mesocosm Validation, and Pilot-Scale Field Trials. Each stage has inputs, processes, outputs, and specific hurdles like ecological components, efficacy assessment, and technical hurdles. The process advances from basic research to deployment, highlighting the transition from promising candidates to full-scale implementation, addressing economic and logistical feasibility.</alt-text>
</graphic>
</fig>
</sec>
</sec>
<sec id="s6-4">
<label>6.4</label>
<title>Translating mechanistic insights into field strategies</title>
<p>The fundamental understanding of degradation mechanisms, as synthesized in <xref ref-type="sec" rid="s4">Sections 4</xref>, <xref ref-type="sec" rid="s5">5</xref>, must directly inform the strategic design and selection of field applications. The choice between <italic>in-situ</italic> and <italic>ex-situ</italic> approaches, and between bioaugmentation and biostimulation, is not merely logistical but is dictated by the inherent biochemical and ecological constraints of the system.</p>
<p>The superior kinetics and specificity of hydrolytic enzymes like PETase for PET depolymerization argue strongly for their deployment in controlled, <italic>ex-situ</italic> bioreactor systems. Here, the optimal temperature, pH, and enzyme-to-substrate ratios required for high-efficiency breakdown (<xref ref-type="sec" rid="s5-2">Section 5.2</xref>) can be maintained, turning waste into valuable monomers for a circular economy (<xref ref-type="sec" rid="s7-1">Section 7.1</xref>). In contrast, the slow, oxidative breakdown of polyolefins (PE, PP) by enzymes like alkane hydroxylases, which often requires abiotic pre-weathering and yields minimal energy gain for microbes, renders large-scale <italic>in-situ</italic> bioremediation of these polymers ecologically and energetically implausible with current technology.</p>
<p>Furthermore, the microbial ecology of the plastisphere (<xref ref-type="sec" rid="s3">Section 3</xref>) underscores why simple bioaugmentation with a single, high-performing strain is likely to fail. The degradation process relies on successional biofilm development and consortia-based metabolism, where primary degraders initiate breakdown and secondary specialists mineralize the intermediates through cross-feeding (<xref ref-type="sec" rid="s3-2">Sections 3.2</xref>, <xref ref-type="sec" rid="s4-2">4.2</xref>). Introducing a single non-native strain ignores this requisite functional network and subjects it to intense competition and predation (<xref ref-type="sec" rid="s6-2">Section 6.2</xref>). Therefore, biostimulation of native plastisphere communities tailored to the polymer type present emerges as a more ecologically sound strategy, as it leverages pre-adapted, complex consortia. Where bioaugmentation is considered, it should involve engineered, ecologically competent consortia designed with synthetic ecology principles to perform coordinated degradation, though their release is mired in the significant regulatory challenges associated with GMOs (<xref ref-type="sec" rid="s7-2-2">Section 7.2.2</xref>). Consequently, the most viable near-term applications are those where biological mechanisms align with engineering and regulatory feasibility: enzymatic recycling in contained facilities for specific polymers like PET, and carefully managed biostimulation for targeted <italic>in-situ</italic> remediation where conditions and polymer types are favorable.</p>
</sec>
</sec>
<sec id="s7">
<label>7</label>
<title>Policy, economic, and biotechnological implications</title>
<p>The scientific understanding of bacterial degradation, as synthesized in previous sections, must be contextualized within broader societal and industrial frameworks to realize its potential.</p>
<sec id="s7-1">
<label>7.1</label>
<title>The role of bacterial degradation in circular economy models for plastics</title>
<p>The current global plastic economy is predominantly linear (take-make-dispose), a model that is environmentally unsustainable. The concept of a circular economy aims to eliminate waste and keep materials in continuous cycles of use. Within this framework, bacterial degradation offers unique value propositions, primarily at the recycling and recovery stages.</p>
<sec id="s7-1-1">
<label>7.1.1</label>
<title>Biological recycling (bio-recycling)</title>
<p>For polymers like PET, enzymatic degradation can depolymerize waste plastic back into its core monomers, terephthalic acid (TPA) and ethylene glycol (EG). These monomers can then be repolymerized into new, virgin-quality PET, creating a closed-loop system that is not possible with conventional mechanical recycling, which downcycles material quality (<xref ref-type="bibr" rid="B24">Joseph et al., 2024</xref>). This process, known as chemobiosis or biological depolymerization, can handle mixed-color or contaminated plastic streams that are challenging for mechanical methods.</p>
</sec>
<sec id="s7-1-2">
<label>7.1.2</label>
<title>Waste valorization</title>
<p>Beyond closed-loop recycling, bacterial processes can be designed to convert plastic waste into value-added products. Through metabolic engineering, bacteria can be tailored to funnel plastic degradation intermediates not into CO<sub>2</sub>, but into building blocks for bioplastics (e.g., PHA), biosurfactants, or other biochemicals, thereby creating an economic incentive for plastic waste collection and processing (<xref ref-type="bibr" rid="B37">Liu Z et al., 2025</xref>). However, this review identifies that bacterial degradation is not a panacea. Its role is complementary to other strategies, most effectively targeting plastic waste that has escaped collection, has been too degraded for mechanical recycling, or is a contaminant in organic waste streams.</p>
</sec>
</sec>
<sec id="s7-2">
<label>7.2</label>
<title>Regulatory frameworks and gaps for bioremediation technologies</title>
<p>The translation of promising bacterial degradation strategies from the laboratory to field-scale environmental applications faces a critical bottleneck: the existing regulatory landscape. Current international frameworks, such as the US Toxic Substances Control Act (TSCA) and the EU&#x2019;s Registration, Evaluation, Authorisation and Restriction of Chemicals (REACH) regulations, are primarily designed to manage the risks of chemical substances. These systems are not well-adapted to assess the complex risks, benefits, and long-term ecological consequences of novel bioremediation agents, particularly for <italic>in-situ</italic> remediation of microplastics. Our analysis identifies three foundational gaps that must be addressed to enable safe and effective deployment.</p>
<sec id="s7-2-1">
<label>7.2.1</label>
<title>Ambiguous definitions and standardized endpoints</title>
<p>A primary obstacle is the lack of universally accepted, environmentally relevant definitions for &#x201c;successful&#x201d; biodegradation. Regulatory approval often hinges on simplistic metrics like mass loss, which does not account for the complete environmental fate of the polymer. A more robust framework must mandate evidence of ultimate mineralization, tracking polymer carbon through to carbon dioxide, water, and biomass, rather than stopping at fragmentation or partial degradation. Furthermore, standardized protocols for assessing the ecotoxicity of degradation by-products are absent. This is a critical oversight, as the breakdown of complex polymers can release monomers, oligomers, and chemical additives (e.g., phthalates, plasticizers) that may be more bioavailable and toxic than the parent material (<xref ref-type="bibr" rid="B20">Hu et al., 2025</xref>; <xref ref-type="bibr" rid="B74">Yadav et al., 2025</xref>). Regulations risk permitting a remediation strategy that merely converts a visible macro- or micro-pollutant into an invisible, but potentially more harmful, chemical mixture, thereby exacerbating ecological toxicity.</p>
</sec>
<sec id="s7-2-2">
<label>7.2.2</label>
<title>The genetically modified organism (GMO) conundrum</title>
<p>The most efficient and engineered degradative strains, whether optimized for enzyme performance, substrate range, or ecological resilience, typically fall under strict GMO regulations. The pathway for the deliberate environmental release of GMOs is predicated on a precautionary principle, making it extremely stringent, costly, and time-consuming. While this caution is valid for biosafety, it often stifles innovation by overlooking a formal risk-benefit analysis specific to pollution remediation. This creates a significant barrier to deploying potentially transformative solutions (<xref ref-type="bibr" rid="B34">Lea-Smith et al., 2025</xref>). The challenge is further complicated by emerging strategies like genetic bioaugmentation, which involves the use of mobile genetic elements (e.g., conjugative plasmids) to transfer degradative genes to indigenous, well-adapted bacterial populations <italic>in situ</italic>. Such approaches blur the line between a contained, introduced GMO and a natural, engineered community, creating a regulatory gray area that existing frameworks are not designed to evaluate.</p>
</sec>
<sec id="s7-2-3">
<label>7.2.3</label>
<title>Regulatory dichotomy: <italic>in-situ</italic> vs. <italic>ex-situ</italic> applications</title>
<p>The regulatory pathway bifurcates sharply based on the context of application, highlighting a more feasible near-term avenue for the technology. For <italic>in situ</italic> applications (direct release into open environments), the regulatory hurdles described above are paramount. Each of the gaps in definition, by-product assessment, and GMO status must be navigated, requiring extensive pre-market environmental risk assessments that are not yet standardized for micro-plastic-degrading biocatalysts. For <italic>ex-situ</italic> applications (contained systems like bioreactors), the regulatory path is markedly clearer and analogous to the well-established regulation of industrial enzymes and fermentation processes. In controlled bioprocessing facilities, risks are contained, monitoring is direct, and the focus shifts to worker safety, product purity, and industrial effluent standards. This distinction underscores that the most viable and immediate application of bacterial and enzymatic degradation lies not in the diffuse cleanup of oceans, but in targeted bio-recycling operations within waste management infrastructure, where plastics can be collected, concentrated, and processed in a controlled, regulated environment.</p>
</sec>
</sec>
<sec id="s7-3">
<label>7.3</label>
<title>Harnessing bacterial enzymes (PETase) for industrial bioprocessing of plastic waste</title>
<p>The most immediate and technologically feasible application of the discoveries in this field is not the <italic>in-situ</italic> cleanup of environmental microplastics, but the development of <italic>ex-situ</italic> enzyme-based biorecycling processes. This approach leverages the specificity and efficiency of bacterial enzymes in controlled industrial settings.</p>
<sec id="s7-3-1">
<label>7.3.1</label>
<title>Enzyme engineering</title>
<p>The native PETase from <italic>I. sakaiensis</italic> has been the starting point for extensive protein engineering efforts. Through rational design and directed evolution, scientists have created mutant enzymes with significantly enhanced thermal stability, catalytic activity, and ability to degrade higher-crystallinity PET, making them more suitable for industrial processes (<xref ref-type="bibr" rid="B71">Wang et al., 2024</xref>).</p>
</sec>
<sec id="s7-3-2">
<label>7.3.2</label>
<title>Process integration</title>
<p>Pilot-scale projects are now demonstrating the integration of engineered enzymes into waste management workflows. For example, post-consumer PET waste is mechanically shredded, pre-treated with heat to reduce crystallinity, and then fed into bioreactors containing the engineered hydrolases. The resulting monomers are purified and sold back to plastic manufacturers (<xref ref-type="bibr" rid="B10">Cui et al., 2024</xref>). This model transforms plastic waste from a liability into a feedstock. <xref ref-type="table" rid="T8">Table 8</xref> compares the potential applications of bacterial degradation, highlighting the stark contrast between the challenges of environmental remediation and the opportunities in industrial bioprocessing, as further conceptualized in <xref ref-type="fig" rid="F6">Figure 6</xref>.</p>
<fig id="F6" position="float">
<label>FIGURE 6</label>
<caption>
<p>The evolving role of bacterial solutions in the plastic value chain.</p>
</caption>
<graphic xlink:href="feart-14-1752600-g006.tif">
<alt-text content-type="machine-generated">Flowchart illustrating the plastic lifecycle from virgin fossil fuel through plastic production, consumer use, and landfill/environment. It highlights enzymatic bioreactor recycling: plastic waste collection, processing, and conversion into value-added biochemicals. Dashed lines indicate downcycled products and in-situ bioremediation as a long-term goal.</alt-text>
</graphic>
</fig>
</sec>
</sec>
</sec>
<sec id="s8">
<label>8</label>
<title>Critical analysis and research gaps</title>
<p>This systematic review has synthesized a rapidly growing body of evidence demonstrating the potential for bacterial degradation of microplastics. However, a critical appraisal of the literature is imperative to contextualize these findings, assess their reliability, and steer future research away from established pitfalls and toward the most pressing unanswered questions.</p>
<sec id="s8-1">
<label>8.1</label>
<title>Assessment of the strength and limitations of the current evidence base</title>
<p>The overall evidence base is characterized by high enthusiasm but variable methodological rigor, with the strength of the field lying in the consistent, independent identification of key bacterial taxa (e.g., <italic>Pseudomonas</italic>, <italic>Bacillus</italic>) across studies, the elucidation of specific enzymatic pathways (e.g., PETase/MHETase), and the clear demonstration of principle that biodegradation is possible; however, significant limitations temper the translation of these findings, including the fact that a vast majority of studies provide proof-of-principle under idealized laboratory conditions that are not representative of environmental realities, and many reported degradation rates, while statistically significant, are ecologically irrelevant on short timescales (e.g., &#x3c;10% weight loss over several months) (<xref ref-type="bibr" rid="B66">Singh and Borthakur, 2018</xref>).</p>
<p>Further limitations arise from the lack of environmental context, as studies frequently use pure bacterial cultures and pristine, often commercially sourced polymer powders or films, neglecting the effects of environmental aging, biofilm succession, and the complex chemical cocktail of additives and adsorbed pollutants present in environmental microplastics (<xref ref-type="bibr" rid="B18">Heris, 2024</xref>; <xref ref-type="bibr" rid="B11">Deng et al., 2024</xref>). There is insufficient analytical rigor, because while many studies use SEM and FTIR, fewer employ the more definitive metrics of mineralization (e.g., respirometry to measure CO<sub>2</sub> evolution) or polymer integrity (e.g., GPC to confirm chain scission), leaving open the possibility that observed changes are surface modifications rather than substantive degradation (<xref ref-type="bibr" rid="B14">Grima et al., 2000</xref>; <xref ref-type="bibr" rid="B26">Keridou et al., 2020</xref>; <xref ref-type="bibr" rid="B65">Silva et al., 2023</xref>; <xref ref-type="bibr" rid="B68">Tarhan and Kestek, 2024</xref>).</p>
</sec>
<sec id="s8-2">
<label>8.2</label>
<title>Identification of high-risk-of-bias areas and methodological heterogeneity</title>
<p>Our analysis identified specific areas where the risk of bias is high and methodological heterogeneity impedes cross-study comparison and meta-analysis.</p>
<sec id="s8-2-1">
<label>8.2.1</label>
<title>High-risk-of-bias areas</title>
<p>This analysis identified several key areas that introduce a high risk of bias in the current literature. A prevalent issue is the lack of adequate sterile controls to account for abiotic degradation, which can lead to an overestimation of biological efficacy. Withal, a likely publication bias exists due to the under-reporting of negative or inconclusive results, skewing the field toward overly optimistic outcomes. Compounding these problems is the frequent failure to fully characterize the starting plastic material, as insufficient documentation of properties like molecular weight, crystallinity, and additive content prevents meaningful correlation between polymer structure and degradability.</p>
</sec>
<sec id="s8-2-2">
<label>8.2.2</label>
<title>Methodological heterogeneity</title>
<p>Methodological heterogeneity is a significant challenge in this field, stemming from a lack of standardized protocols. Critical variables, such as the polymer form (film, powder, pellet), inoculum source and preparation (pure culture vs. enrichment culture), incubation conditions (media, temperature, shaking speed), and the duration of experiments, differ vastly across studies. This considerable variability currently precludes a meaningful quantitative meta-analysis and highlights the urgent necessity for standardized testing guidelines (see recommended research directions in <xref ref-type="table" rid="T9">Table 9</xref>).</p>
</sec>
</sec>
<sec id="s8-3">
<label>8.3</label>
<title>Key knowledge gaps: long-term efficacy, ecotoxicity of by-products, and engineered consortia</title>
<p>Beyond methodological issues, this review identifies several fundamental scientific gaps that represent the Frontier of research in this field.</p>
<sec id="s8-3-1">
<label>8.3.1</label>
<title>Long-term efficacy and environmental fate</title>
<p>There is a near-total absence of data on the long-term (multi-year) persistence and activity of plastic-degrading bacteria in the environment. It is unknown whether degradation follows a linear progression, plateaus, or if the microbial community evolves to become more or less efficient over time. Crucially, the ultimate fate of the plastic carbon remains largely uncharacterized. While mineralization to CO<sub>2</sub> is the desired endpoint, it is essential to determine what fraction of the carbon is instead incorporated into microbial biomass, transformed into persistent dissolved organic carbon, or sequestered in an intermediate, partially degraded state (<xref ref-type="bibr" rid="B31">Krause et al., 2020</xref>; <xref ref-type="bibr" rid="B18">Heris, 2024</xref>). Without this understanding, the true environmental impact and carbon footprint of bioremediation cannot be assessed.</p>
</sec>
<sec id="s8-3-2">
<label>8.3.2</label>
<title>Ecotoxicity of by-products: a critical oversight</title>
<p>A profound and frequently overlooked gap concerns the potential toxicity of intermediary degradation products, as the current literature predominantly celebrates the disappearance of the plastic particle but fails to systematically assess the biological impact of the resulting chemical mixture, which is a critical omission, since bioremediation risks solving one problem (MPs) while exacerbating another (chemical pollution), and therefore future studies must move beyond simple degradation metrics to mandate integrated ecotoxicological assessments.</p>
<p>For different polymers, degradation can release a suite of potentially hazardous chemicals, including phthalates and bisphenol A (BPA) from PVC and polycarbonates, styrene oligomers and phenylacetaldehyde from polystyrene (PS) degradation, and terephthalic acid (TPA) and ethylene glycol from PET at ecologically relevant concentrations, and many of these compounds are known endocrine disruptors or have demonstrated toxicity to aquatic organisms.</p>
<p>A robust framework for monitoring and assessment is needed, combining targeted chemical analysis (e.g., LC-MS/MS for specific monomers and additives) with whole-mixture bioassays using sensitive indicator species across trophic levels (e.g., algae, daphnids, fish embryos), as this dual approach is necessary to identify both known culprits and unforeseen synergistic toxic effects.</p>
</sec>
<sec id="s8-3-3">
<label>8.3.3</label>
<title>Rational design and ecology of engineered consortia</title>
<p>While consortia are consistently shown to be more effective than single isolates (<xref ref-type="sec" rid="s4-2">Section 4.2</xref>), their development remains largely <italic>ad hoc</italic>, relying on random enrichment cultures. A significant gap exists in the application of synthetic ecology principles to rationally design stable, cooperative, and effective microbial communities for plastic degradation.</p>
<sec id="s8-3-3-1">
<label>8.3.3.1</label>
<title>Guiding principles for design</title>
<p>Future efforts should focus on constructing consortia based on complementary metabolic pathways, ensuring that the breakdown products from one member (e.g., PET oligomers from an <italic>Ideonella</italic> strain) serve as the optimal substrate for another (e.g., a specialist in TPA metabolism). Stability can be engineered through cross-feeding dependencies and quorum sensing networks to regulate population dynamics and enzyme production. Furthermore, functional redundancy (including multiple species capable of key steps) should be built in to confer resilience against environmental fluctuations.</p>
</sec>
<sec id="s8-3-3-2">
<label>8.3.3.2</label>
<title>Environmental integration and safety</title>
<p>Research must also address how to ensure the ecological competence and containment of designed consortia. This includes understanding their interactions with indigenous microbes, their potential for horizontal gene transfer, and strategies for their eventual removal or die-off after remediation is complete, ensuring they do not become persistent, disruptive elements in the ecosystem (<xref ref-type="bibr" rid="B22">Johns et al., 2016</xref>). These critical safety and integration challenges represent key priorities on the strategic research roadmap for advancing bio-based remediation from the laboratory to the field, as visualized in <xref ref-type="fig" rid="F7">Figure 7</xref>.</p>
<fig id="F7" position="float">
<label>FIGURE 7</label>
<caption>
<p>Bridging the knowledge gap: a strategic roadmap for future research.</p>
</caption>
<graphic xlink:href="feart-14-1752600-g007.tif">
<alt-text content-type="machine-generated">Flowchart depicting &#x22;Standardized Methodologies &#x26; Data Reporting&#x22; at the center, with arrows pointing to four sections: &#x22;Environmental Realism&#x22; with a beaker illustration, &#x22;Mechanism &#x26; Prediction&#x22; with DNA and microchip graphics, &#x22;Application &#x26; Risk Assessment&#x22; with gears labeled &#x22;Engineered Consortia,&#x22; and &#x22;Eco-Toxicity Screening&#x22; with a fish and test tube. Each section poses questions related to environmental degradation and assessment.</alt-text>
</graphic>
</fig>
</sec>
</sec>
</sec>
</sec>
<sec sec-type="conclusion" id="s9">
<label>9</label>
<title>Conclusion</title>
<p>This systematic review, synthesizing evidence from 80 studies, unequivocally demonstrates that bacterial degradation represents a scientifically valid and promising pathway for addressing the global challenge of microplastic pollution. The findings confirm that a phylogenetically diverse consortium of bacteria, primarily within the genera <italic>Pseudomonas</italic>, <italic>Bacillus</italic>, and <italic>Rhodococcus</italic>, possesses the enzymatic machinery including hydrolases, oxidoreductases, and laccases to initiate the breakdown of pervasive polymers like PET, PE, PP, and PS. The process is an active, regulated metabolic response, often enhanced within the structured, synergistic environment of the plastisphere biofilm.</p>
<p>However, the transition from laboratory promise to environmental panacea is fraught with significant challenges. A central conclusion of this review is the stark disparity between degradation efficacy in controlled settings and the realities of complex aquatic environments. While PET shows remarkable susceptibility to enzymatic hydrolysis, the recalcitrance of polyolefins (PE, PP) remains a major bottleneck, with reported degradation rates being slow and often ecologically insignificant on short timescales. The critical analysis further reveals that the field is hampered by methodological heterogeneity, a lack of environmental context in many studies, and an insufficient focus on the ecotoxicological risks of degradation by-products.</p>
<p>Therefore, the most viable and immediate application of this technology lies not in the diffuse cleanup of open waters, but in targeted, <italic>ex-situ</italic> systems. The enzymatic biorecycling of PET into its constituent monomers for a circular economy stands out as a technologically mature and economically incentivized pathway. For <italic>in-situ</italic> remediation, a hybrid bioaugmentation and biostimulation approach, tailored to specific polymer pollutants, holds potential but requires breakthroughs in overcoming ecological barriers, monitoring efficacy, and navigating regulatory frameworks.</p>
<p>Future research must pivot towards closing the critical gaps identified in this review. Priorities include conducting long-term, environmentally realistic mesocosm studies, implementing standardized protocols to enable cross-study comparisons, and mandating integrated ecotoxicological assessments. The rational design of stable, effective microbial consortia and the development of robust tools for <italic>in-situ</italic> monitoring are essential next steps. By embracing this strategic, interdisciplinary, and critical pathway, the scientific community can translate the compelling potential of plastic-degrading bacteria into tangible, responsible, and effective solutions for mitigating plastic pollution.</p>
</sec>
</body>
<back>
<sec sec-type="data-availability" id="s10">
<title>Data availability statement</title>
<p>The original contributions presented in the study are included in the article/<xref ref-type="sec" rid="s16">Supplementary Material</xref>, further inquiries can be directed to the corresponding author.</p>
</sec>
<sec sec-type="author-contributions" id="s11">
<title>Author contributions</title>
<p>MB: Writing &#x2013; original draft, Writing &#x2013; review and editing. UA: Writing &#x2013; review and editing, Writing &#x2013; original draft. TT: Writing &#x2013; review and editing, Writing &#x2013; original draft.</p>
</sec>
<sec sec-type="COI-statement" id="s13">
<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="s14">
<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="s15">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<sec sec-type="supplementary-material" id="s16">
<title>Supplementary material</title>
<p>The Supplementary Material for this article can be found online at: <ext-link ext-link-type="uri" xlink:href="https://www.frontiersin.org/articles/10.3389/feart.2026.1752600/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/feart.2026.1752600/full&#x23;supplementary-material</ext-link>
</p>
<supplementary-material xlink:href="Table1.docx" id="SM1" mimetype="application/docx" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</sec>
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</fn>
<fn fn-type="custom" custom-type="reviewed-by">
<p>
<bold>Reviewed by:</bold> <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/2270497/overview">imran IqbaL</ext-link>, Yale University, United States</p>
<p>
<ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/3308317/overview">Milena Roberta Freire Da Silva</ext-link>, Federal University of Pernambuco, Brazil</p>
</fn>
</fn-group>
</back>
</article>