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<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Microbiol.</journal-id>
<journal-title>Frontiers in Microbiology</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Microbiol.</abbrev-journal-title>
<issn pub-type="epub">1664-302X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fmicb.2024.1521048</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Microbiology</subject>
<subj-group>
<subject>Mini Review</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Fish gut microbiome and its application in aquaculture and biological conservation</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name><surname>Kanika</surname> <given-names>Nusrat Hasan</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
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<name><surname>Liaqat</surname> <given-names>Nusrat</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff4"><sup>4</sup></xref>
<xref ref-type="aff" rid="aff5"><sup>5</sup></xref>
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<contrib contrib-type="author">
<name><surname>Chen</surname> <given-names>Huifan</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
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<name><surname>Ke</surname> <given-names>Jing</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
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<name><surname>Lu</surname> <given-names>Guoqing</given-names></name>
<xref ref-type="aff" rid="aff6"><sup>6</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>&#x0002A;</sup></xref>
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<name><surname>Wang</surname> <given-names>Jun</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
<xref ref-type="corresp" rid="c002"><sup>&#x0002A;</sup></xref>
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<name><surname>Wang</surname> <given-names>Chenghui</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
<xref ref-type="corresp" rid="c003"><sup>&#x0002A;</sup></xref>
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<aff id="aff1"><sup>1</sup><institution>Key Laboratory of Freshwater Aquatic Genetic Resources, Ministry of Agriculture and Rural Affairs, Shanghai Ocean University</institution>, <addr-line>Shanghai</addr-line>, <country>China</country></aff>
<aff id="aff2"><sup>2</sup><institution>National Demonstration Center for Experimental Fisheries Science Education, Shanghai Ocean University</institution>, <addr-line>Shanghai</addr-line>, <country>China</country></aff>
<aff id="aff3"><sup>3</sup><institution>Shanghai Engineering Research Center of Aquaculture, Shanghai Ocean University</institution>, <addr-line>Shanghai</addr-line>, <country>China</country></aff>
<aff id="aff4"><sup>4</sup><institution>Shanghai Collaborative Innovation for Aquatic Animal Genetics and Breeding, Shanghai Ocean University</institution>, <addr-line>Shanghai</addr-line>, <country>China</country></aff>
<aff id="aff5"><sup>5</sup><institution>National Experimental Teaching Demonstration Centre for Aquatic Sciences, Shanghai Ocean University</institution>, <addr-line>Shanghai</addr-line>, <country>China</country></aff>
<aff id="aff6"><sup>6</sup><institution>Department of Biology, University of Nebraska at Omaha</institution>, <addr-line>Omaha, NE</addr-line>, <country>United States</country></aff>
<author-notes>
<fn fn-type="edited-by"><p>Edited by: Lifeng Zhu, Nanjing University of Chinese Medicine, China</p></fn>
<fn fn-type="edited-by"><p>Reviewed by: Yaqiu Liu, Chinese Academy of Fishery Sciences, China</p>
<p>Wancai Xia, China West Normal University, China</p></fn>
<corresp id="c001">&#x0002A;Correspondence: Guoqing Lu <email>glu3&#x00040;unomaha.edu</email></corresp>
<corresp id="c002">Jun Wang <email>wangjun&#x00040;shou.edu.cn</email></corresp>
<corresp id="c003">Chenghui Wang <email>wangch&#x00040;shou.edu.cn</email></corresp>
</author-notes>
<pub-date pub-type="epub">
<day>07</day>
<month>01</month>
<year>2025</year>
</pub-date>
<pub-date pub-type="collection">
<year>2024</year>
</pub-date>
<volume>15</volume>
<elocation-id>1521048</elocation-id>
<history>
<date date-type="received">
<day>01</day>
<month>11</month>
<year>2024</year>
</date>
<date date-type="accepted">
<day>09</day>
<month>12</month>
<year>2024</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x000A9; 2025 Kanika, Liaqat, Chen, Ke, Lu, Wang and Wang.</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>Kanika, Liaqat, Chen, Ke, Lu, Wang and Wang</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/"><p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p></license>
</permissions>
<abstract>
<p>Understanding the diversity and function of fish gut microbiomes has advanced substantially, yet many aspects remain poorly understood, particularly the interplay among microbiota, host species, and environmental factors in the context of conservation. This review explores the composition and abundance of gut bacterial communities in key aquaculture fish groups&#x02014;cyprinids, ictalurids (catfish), salmonids, and cichlids (tilapia)&#x02014;alongside the model organism zebrafish, across diverse geographic regions. The findings highlight environmental habitats and host species as primary determinants of gut microbiome structure, offering a global perspective on these microbial communities. Across all fish groups, the phyla Firmicutes, Fusobacteria, and Proteobacteria consistently dominated, while temperate, sub-equatorial, and sub-tropical regions exhibited the highest microbiome diversity, underscoring the contribution of taxonomic and environmental factors. The gut bacterial diversity of farm-raised fish shows a significant divergence from that of wild-caught fish, reflecting the impacts of ecological and management differences. Understanding the dynamic responses of fish gut microbiota is vital for guiding conservation efforts, safeguarding aquatic biodiversity, and advancing sustainable aquaculture practices. Future research should leverage innovative techniques and integrative approaches, both experimental and theoretical, to uncover the functional roles of microbiomes and predict their responses to environmental changes. Expanding geographic and taxonomic coverage will be critical for creating a comprehensive framework to inform global aquaculture and conservation strategies. Collectively, this perspective highlights the transformative potential of microbiome research in addressing global challenges in aquaculture and conservation biology.</p></abstract>
<kwd-group>
<kwd>fish gut microbiome</kwd>
<kwd>aquaculture species</kwd>
<kwd>microbial diversity</kwd>
<kwd>host association</kwd>
<kwd>environment impact</kwd>
<kwd>conservation biology</kwd>
</kwd-group>
<counts>
<fig-count count="1"/>
<table-count count="1"/>
<equation-count count="0"/>
<ref-count count="106"/>
<page-count count="13"/>
<word-count count="8287"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Microbial Symbioses</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec sec-type="intro" id="s1">
<title>Introduction</title>
<p>Microbiomes play critical roles in maintaining ecosystem health, nutrient cycling, and climate regulation (Lennon et al., <xref ref-type="bibr" rid="B40">2023</xref>). Within aquatic environments, fish, the most diverse group of vertebrates, are host to complex microbial communities that significantly impact their physiology and the health of the surrounding environment (Lorgen-Ritchie et al., <xref ref-type="bibr" rid="B52">2023</xref>). The fish gut microbiome presents an invaluable window into host-microbiota-environment interactions, offering insights with direct implications for aquaculture and conservation. The composition of fish gut microbiota is shaped by a combination of environmental, biological, and behavioral factors, which collectively influence microbial communities across diverse fish species (Nayfach et al., <xref ref-type="bibr" rid="B64">2021</xref>; Bertoncin et al., <xref ref-type="bibr" rid="B8">2022</xref>). These microbiomes are sensitive to environmental conditions, including water temperature, oxygen levels, pH, and salinity, as well as intrinsic factors such as feeding behaviors, life stages, and various anthropogenic influences (Liu et al., <xref ref-type="bibr" rid="B49">2016</xref>; Li et al., <xref ref-type="bibr" rid="B43">2017a</xref>; Du et al., <xref ref-type="bibr" rid="B20">2019</xref>; Zotta et al., <xref ref-type="bibr" rid="B106">2019</xref>; Huang et al., <xref ref-type="bibr" rid="B31">2020</xref>; Mukherjee et al., <xref ref-type="bibr" rid="B63">2020</xref>). The dynamic interplay between fish and their gut microbiota plays a crucial role in shaping fish communities, making this an important area of study for better understanding and managing aquatic biodiversity.</p>
<p>Beyond contributing to essential ecosystem functions, such as nutrient cycling and climate regulation, gut microbiota are also critical for individual health. Alterations in microbial communities can affect phenotypic traits, immune mechanisms, and animal fitness in response to climate change, as physiological functions originate mostly from the gut (Dinan and Cryan, <xref ref-type="bibr" rid="B18">2016</xref>; Mohajeri et al., <xref ref-type="bibr" rid="B60">2018</xref>; Sepulveda and Moeller, <xref ref-type="bibr" rid="B74">2020</xref>). Microbiomes are leveraged to enhance fatty acid production in muscle tissue and improve fish development (Eichmiller et al., <xref ref-type="bibr" rid="B22">2016</xref>; Stephens et al., <xref ref-type="bibr" rid="B81">2016</xref>; Mohajeri et al., <xref ref-type="bibr" rid="B60">2018</xref>; Aldars-Garc&#x000ED;a et al., <xref ref-type="bibr" rid="B2">2021</xref>; Asnicar et al., <xref ref-type="bibr" rid="B4">2021</xref>; Chen et al., <xref ref-type="bibr" rid="B11">2021</xref>; Zhang et al., <xref ref-type="bibr" rid="B100">2022</xref>; Yin et al., <xref ref-type="bibr" rid="B98">2023</xref>). The gut microbiome helps protect the intestinal barrier, prevent the overgrowth of opportunistic pathogens, and modulate the host immune system, all of which are crucial for maintaining fish health (Merrifield and Rodiles, <xref ref-type="bibr" rid="B59">2015</xref>; Llewellyn et al., <xref ref-type="bibr" rid="B51">2016</xref>; Nohesara et al., <xref ref-type="bibr" rid="B65">2023</xref>). Conversely, disruption of the microbial balance can result in the proliferation of harmful bacteria, leading to disease outbreaks in aquaculture settings (Talwar et al., <xref ref-type="bibr" rid="B83">2018</xref>; Vargas-albores et al., <xref ref-type="bibr" rid="B87">2021</xref>; Wang et al., <xref ref-type="bibr" rid="B88">2021</xref>). Therefore, understanding and manipulating the fish gut microbiome has become an important strategy for developing sustainable and disease-resistant aquaculture practices.</p>
<p>This review investigates how microbial abundance varies in response to temperature, habitat, and taxonomic differences across major fish groups in global aquaculture. We focus on cyprinids, ictalurids (catfish), salmonids, and cichlids (tilapia), which are economically significant aquaculture species (Lu and Luo, <xref ref-type="bibr" rid="B54">2020</xref>). Additionally, we compare the gut microbiomes of farmed fish, which are raised in controlled environments with standardized diets, with those of wild-caught fish, which interact directly with their natural habitats, to understand how these contrasting conditions influence fish gut microbiomes. This comparative approach will help identify key environmental and dietary factors shaping the gut microbiome and highlight bacterial groups particularly sensitive to these variables. By highlighting these variations and their underlying causes, this review offers valuable insights into the role of the gut microbiome in promoting fish resilience and health under changing environmental conditions. These insights are essential for informing conservation strategies and optimizing sustainable aquaculture practices worldwide.</p></sec>
<sec id="s2">
<title>Fish gut microbiomes varying across habitats, climatic zones, and feeding behaviors</title>
<p>Gut microbiota composition varies among fish taxa, with hosts from the same taxonomic group generally exhibiting more similar gut microbiota than those from different groups; however, biological factors such as feeding habits can lead to remarkable differences within taxa (Huang et al., <xref ref-type="bibr" rid="B31">2020</xref>). Distinct gut microbiome compositions were observed across different fish groups, with Proteobacteria, Fusobacteria, and Firmicutes being the most prevalent (<xref ref-type="fig" rid="F1">Figures 1A</xref>, <xref ref-type="fig" rid="F1">B</xref>). While Actinobacteria was present in cyprinids, salmonids, cichlids, and zebrafish, it was not reported in the catfish group (<xref ref-type="fig" rid="F1">Figure 1B</xref>). Fish species exhibit distinct feeding behaviors across water layers, which shape their gut microbiome composition. Cyprinids, ictalurids (catfish), salmonids, cichlids, and zebrafish, ranging from bottom dwellers to surface feeders, display microbiota variations based on diet and ecological niches (Ang and Petrell, <xref ref-type="bibr" rid="B3">1998</xref>; Rahman et al., <xref ref-type="bibr" rid="B67">2008</xref>; Ramesh and Kiran, <xref ref-type="bibr" rid="B68">2016</xref>; Thomas and Opeh, <xref ref-type="bibr" rid="B84">2018</xref>). The variation in microbiota phyla highlights the impact of feeding behaviors on gut microbiome diversity (<xref ref-type="fig" rid="F1">Figures 1B</xref>, <xref ref-type="fig" rid="F1">C</xref>), emphasizing ecological adaptation (Sinha and Jones, <xref ref-type="bibr" rid="B76">1967</xref>; Magoulick and Lewis, <xref ref-type="bibr" rid="B57">2002</xref>; Watzin et al., <xref ref-type="bibr" rid="B90">2008</xref>).</p>
<fig id="F1" position="float">
<label>Figure 1</label>
<caption><p>Gut microbiome diversity in four fish groups, farm-raised vs. wild-caught, and across climatic zones. <bold>(A)</bold> Fish groups and model species: four key fish groups commonly used in aquaculture&#x02014;cyprinids, ictalurids (catfish), salmonids, and cichlids (tilapia)&#x02014;are depicted, along with the model organism zebrafish. <bold>(B)</bold> Gut microbiome composition across different fish groups: phylogenetic relationships among the fish species are shown on the right, alongside dominant bacterial phyla associated with each fish species shown on the left. <bold>(C)</bold> Microbiome differences between farmed-raised and wild-caught fish: a bar chart illustrates the proportion of each bacterial phylum (represented by bar widths), with bar colors indicating specific fish groups. <bold>(D)</bold> Sampling sites and climate zones: sampling locations from reviewed studies are mapped globally, with climate zones differentiated by distinct colors. <bold>(E)</bold> Dominant bacterial phyla and climatic associations: dominant bacterial phyla in various fish groups are shown in relation to climate zones, with the same color scheme as <bold>(D)</bold>.</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmicb-15-1521048-g0001.tif"/>
</fig>
<p>Studies also revealed the composition of fish gut microbiomes varies based on habitat characteristics and geomorphology, with factors like salinity and the differences between nearshore littoral and offshore profundal zones significantly influencing microbial diversity (Zotta et al., <xref ref-type="bibr" rid="B106">2019</xref>; Huang et al., <xref ref-type="bibr" rid="B31">2020</xref>; Sylvain et al., <xref ref-type="bibr" rid="B82">2020</xref>; Kim et al., <xref ref-type="bibr" rid="B35">2021</xref>; Shang et al., <xref ref-type="bibr" rid="B75">2021</xref>). Farm-raised fishes in controlled environments show higher microbiome abundance, especially of Firmicutes, Fusobacteria, and Proteobacteria, compared to wild-caught fishes, where Firmicutes and Proteobacteria are most abundant (<xref ref-type="fig" rid="F1">Figure 1C</xref>, <xref ref-type="table" rid="T1">Table 1</xref>).</p>
<table-wrap position="float" id="T1">
<label>Table 1</label>
<caption><p>Microbial community composition and abundance in zebrafish, salmonidae, ictalurids (catfish), cichlids (tilapia), and cyprinids across various geographic and climate zones.</p></caption>
<table frame="box" rules="all">
<thead>
<tr style="background-color:#919498;color:#ffffff">
<th valign="top" align="left"><bold>Host</bold></th>
<th valign="top" align="left"><bold>Habitat</bold></th>
<th valign="top" align="left"><bold>Location</bold></th>
<th valign="top" align="left"><bold>Climate zone</bold></th>
<th valign="top" align="left"><bold>Dominant gut microbiota (phylum with <italic>genus</italic> or abundance)</bold></th>
<th valign="top" align="left"><bold>References</bold></th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left"><bold>Zebrafish</bold> <italic>Danio rerio</italic></td>
<td valign="top" align="left">Laboratory</td>
<td valign="top" align="left">Eugene, USA</td>
<td valign="top" align="left">Temperate</td>
<td valign="top" align="left"><bold>Actinobacteriota</bold><italic>: Conexibacter, Mycobacterium, Pseudonocardia</italic>, <bold>Chlamydiota</bold><italic>: Neochlamydia, Parachlamydia</italic>, <bold>Fusobacteriota</bold><italic>: Cetobacterium</italic>, <bold>Proteobacteria</bold><italic>: Aeromonas, Catellibacterium, Comamonas, Delftia, Hyphomicrobium, Pelomonas, Pseudomonas, Raoultella, Shewanella, Sphingomonas, Stenotrophomonas, Thioprofundum, Vibrio, Yersinia</italic></td>
<td valign="top" align="left">Stagaman et al., <xref ref-type="bibr" rid="B79">2017</xref></td>
</tr>
 <tr>
<td/>
<td/>
<td valign="top" align="left">Quebec City, Canada</td>
<td valign="top" align="left">Temperate</td>
<td valign="top" align="left"><bold>Actinobacteriota</bold>: Abundant, <bold>Firmicutes</bold><italic>:</italic> Abundant, <bold>Fusobacteriota</bold>: Abundant, <bold>Proteobacteria</bold>: Mostly Abundant</td>
<td valign="top" align="left">Cornuault et al., <xref ref-type="bibr" rid="B14">2022</xref></td>
</tr>
 <tr>
<td/>
<td/>
<td valign="top" align="left">Eugene, OR, USA</td>
<td valign="top" align="left">Sub-tropical</td>
<td valign="top" align="left"><bold>Firmicutes</bold><italic>: Streptococcus</italic>, <bold>Fusobacteriota</bold>: <italic>Cetobacterium</italic>, <bold>Proteobacteria</bold>: <italic>Aeromonas, Shewanella, Enterobacteriaceae, Diaphorobacter, Pseudomonas, Stenotrophomonas, Vibrio</italic></td>
<td valign="top" align="left">Stephens et al., <xref ref-type="bibr" rid="B81">2016</xref></td>
</tr>
 <tr>
<td/>
<td/>
<td valign="top" align="left">Shanghai, China</td>
<td valign="top" align="left">Sub-tropical</td>
<td valign="top" align="left"><bold>Firmicutes</bold><italic>:</italic> Abundant <bold>Fusobacteriota</bold><italic>:</italic> Mostly Abundant</td>
<td valign="top" align="left">Wang et al., <xref ref-type="bibr" rid="B88">2021</xref></td>
</tr>
 <tr>
<td/>
<td/>
<td valign="top" align="left">Birmingham (UAB), USA</td>
<td valign="top" align="left">Sub-tropical</td>
<td valign="top" align="left"><bold>Actinobacteriota</bold><italic>: Bifidobacterium</italic>, <bold>Firmicutes</bold><italic>: Oscillospira, Ruminococcus, Anaeroglobus</italic>, <bold>Proteobacteria</bold><italic>: Pseudoxanthomonas, Legionella anisa, Legionella norrlandica</italic>, <bold>Verrucomicrobiota</bold>: <italic>Luteolibacter</italic></td>
<td valign="top" align="left">Koo et al., <xref ref-type="bibr" rid="B37">2017</xref></td>
</tr> <tr style="background-color:#dee1e1">
<td valign="top" align="left" colspan="6"><bold>Salmonids</bold></td>
</tr> <tr>
<td valign="top" align="left">Atlantic salmon (<italic>Salmo salar</italic>)</td>
<td valign="top" align="left">Wild caught</td>
<td valign="top" align="left">Jonesboro, USA</td>
<td valign="top" align="left">Temperate</td>
<td valign="top" align="left"><bold>Firmicutes</bold><italic>: Peptostreptococcus, Streptococcus, Peptoniphilus, Gallicola, Peptococcus, Staphylococcus, Candidatus_Bacilloplasma, Bacillus, Shewanella</italic> <bold>Proteobacteria</bold>: <italic>Deefgea, Methylobacterium: Methylorubrum, Holosporaceae, Aeromonas</italic></td>
<td valign="top" align="left">Kara et al., <xref ref-type="bibr" rid="B34">2021</xref></td>
</tr> <tr>
<td valign="top" align="left">Atlantic salmon (<italic>Salmo salar</italic>)</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">Dover, Australia;</td>
<td valign="top" align="left">Temperate</td>
<td valign="top" align="left"><bold>Bacteroidota</bold><italic>: Flavobacteriia, Cloacibacterium</italic> <bold>Firmicutes</bold><italic>: Clostridia, Vibrionaceae, Roseobacter, Bacillus, Aeribacillus, Anoxybacillus Geobacillus</italic>, <bold>Proteobacteria</bold><italic>: Vibrionaceae, Methylobacteriaceae</italic></td>
<td valign="top" align="left">Zarkasi et al., <xref ref-type="bibr" rid="B99">2016</xref></td>
</tr> <tr>
<td valign="top" align="left">Atlantic salmon (<italic>Salmo salar</italic>)</td>
<td valign="top" align="left">Wild caught</td>
<td valign="top" align="left">Hirtshals, Denmark</td>
<td valign="top" align="left">Temperate</td>
<td valign="top" align="left"><bold>Actinobacteriota</bold><italic>: Arthrobacter, Brevibacterium</italic> <bold>Firmicutes</bold>: <italic>Bacillus, Weissella, Leuconostoc, Lactobacillus, Pediococcus, Sporosarcina, Jeotgalicoccus, Streptococcus, Carnobacterium, Lactococcus, Ureibacillus, Geobacillus, Streptococcus</italic> <bold>Proteobacteria</bold><italic>: Erwinia, Sphingomonas, Pseudomonadales</italic></td>
<td valign="top" align="left">Gajardo et al., <xref ref-type="bibr" rid="B24">2016</xref></td>
</tr> <tr>
<td valign="top" align="left">Atlantic salmon (<italic>Salmo salar</italic>)</td>
<td valign="top" align="left">Wild caught</td>
<td valign="top" align="left">Eastern Canada and Western Ireland</td>
<td valign="top" align="left">Temperate</td>
<td valign="top" align="left"><bold>Actinobacteriota</bold>: Abundant <bold>Bacteroidota</bold>: Abundant <bold>Firmicutes</bold>: <italic>Mycoplasma</italic></td>
<td valign="top" align="left">Llewellyn et al., <xref ref-type="bibr" rid="B51">2016</xref></td>
</tr> <tr>
<td valign="top" align="left">Atlantic salmon (<italic>Salmo salar</italic>)</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">Aberdeen, UK</td>
<td valign="top" align="left">Temperate</td>
<td valign="top" align="left"><bold>Bacteroidota</bold>: Abundant <bold>Firmicutes</bold>: Abundant <bold>Proteobacteria</bold>: Abundant <bold>Tenericutes</bold>: Abundant</td>
<td valign="top" align="left">Dehler et al., <xref ref-type="bibr" rid="B17">2017</xref></td>
</tr> <tr>
<td valign="top" align="left">Rainbow trout (<italic>Oncorhynchus mykiss</italic>)</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">B&#x000FC;sum, Germany</td>
<td valign="top" align="left">Temperate</td>
<td valign="top" align="left"><bold>Bacteroidota</bold>: <italic>Bacteroides, Porphyromonas</italic>, <bold>Firmicutes</bold><italic>: Staphylococcus, Vagococcus, Streptococcaceae, Lactobacillus, Lactococcus, Staphylococcus, Streptococcus</italic> <bold>Fusobacteriota</bold><italic>: Fusobacterium, Psychrilyobacter, Fusobacteriaceae</italic>, <bold>Proteobacteria</bold><italic>: Burkholderia, Aliivibrio fischeri, Acinetobacter johnsonii, Moritella, Photobacterium, Pseudoalteromonas, Shewanellaceae, Acinetobacter rhizosphaerae</italic></td>
<td valign="top" align="left">Ratten et al., <xref ref-type="bibr" rid="B69">2017</xref></td>
</tr> <tr>
<td valign="top" align="left">Rainbow trout (<italic>Oncorhynchus mykiss</italic>)</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">Argyll, UK</td>
<td valign="top" align="left">Temperate</td>
<td valign="top" align="left"><bold>Firmicutes</bold><italic>: Lactobacillus, Acetanaerobacterium, Catellicoccus, Streptococcus, Weissella, Leuconostoc, Lactococcus, Enterococcus, Bacillus</italic> <bold>Proteobacteria</bold><italic>: Photobacterium, Pseudomonas, Acinetobacter, Maricurvus, Moritella, Pantoea</italic></td>
<td valign="top" align="left">Lyons et al., <xref ref-type="bibr" rid="B56">2017</xref></td>
</tr> <tr>
<td valign="top" align="left">Atlantic salmon (<italic>Salmo salar</italic>)</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">Washington, USA</td>
<td valign="top" align="left">Sub-tropical</td>
<td valign="top" align="left"><bold>Actinobacteriota</bold><italic>: Actinomycetales</italic> <bold>Bacteroidota</bold><italic>: Sphingobacteriales, Flavobacteriales</italic>, <bold>Firmicutes</bold>: <italic>Lactobacillales</italic>, <bold>Nitrospirota</bold>: <italic>Nitrospirales</italic>, <bold>Proteobacteria</bold><italic>: Aeromonadales, Burkholderiales, Neisseriales, Aeromonas, Shewanella, Rickettsiales</italic></td>
<td valign="top" align="left">Schmidt et al., <xref ref-type="bibr" rid="B73">2016</xref></td>
</tr> <tr>
<td valign="top" align="left">Brook trout (<italic>Salvelinus fontinalis</italic>)</td>
<td valign="top" align="left">Wild caught</td>
<td valign="top" align="left">North-western Italy, Italy</td>
<td valign="top" align="left">Sub-tropical</td>
<td valign="top" align="left"><bold>Actinobacteriota</bold>: Abundant <bold>Firmicutes</bold>: <italic>Bacilli</italic> <bold>Fusobacteriota</bold>: Abundant, <bold>Proteobacteria</bold>: Abundant</td>
<td valign="top" align="left">Mugetti et al., <xref ref-type="bibr" rid="B61">2023</xref></td>
</tr> <tr style="background-color:#dee1e1">
<td valign="top" align="left" colspan="6"><bold>Ictalurids (catfish)</bold></td>
</tr> <tr>
<td valign="top" align="left">Catfish (<italic>Ictalurus punctatus</italic>)</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">Ferndale, USA</td>
<td valign="top" align="left">Sub-tropical</td>
<td valign="top" align="left"><bold>Firmicutes</bold><italic>: Streptococcus, Lactobacillus</italic> <bold>Fusobacteriota</bold><italic>: Cetobacterium</italic> <bold>Proteobacteria</bold><italic>: Bradyrhizobium, Plesiomonas, Comamonadaceae, Enterobacteriaceae, Bradyrhizobium</italic></td>
<td valign="top" align="left">Bledsoe et al., <xref ref-type="bibr" rid="B9">2016</xref></td>
</tr> <tr>
<td valign="top" align="left">Southern catfish (<italic>Silurus meridionalis</italic>)</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">Wuhan, China</td>
<td valign="top" align="left">Sub-equatorial</td>
<td valign="top" align="left"><bold>Bacteroidota</bold>: Abundant <bold>Fusobacteriota</bold>: Abundant <bold>Proteobacteria</bold><italic>: Enterobacteriaceae, Plesiomonas, unclassified Aeromonadaceae, Morganella</italic> <bold>Tenericutes</bold>: Abundant</td>
<td valign="top" align="left">Zhang et al., <xref ref-type="bibr" rid="B102">2018</xref></td>
</tr> <tr>
<td valign="top" align="left">Southern catfish, <italic>Silurus meridionalis</italic></td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">Chongqing, China</td>
<td valign="top" align="left">Sub-equatorial</td>
<td valign="top" align="left"><bold>Firmicutes</bold><italic>: Clostridiaceae, Clostridium, Bacillus</italic> <bold>Fusobacteriota</bold><italic>: Cetobacterium</italic> <bold>Proteobacteria</bold><italic>: Plesiomonas</italic></td>
<td valign="top" align="left">Zhang et al., <xref ref-type="bibr" rid="B101">2017</xref></td>
</tr> <tr>
<td valign="top" align="left">Yellow catfish (<italic>Pelteobagrus fulvidraco</italic>)</td>
<td valign="top" align="left">Wild caught</td>
<td valign="top" align="left">Wuhan, China</td>
<td valign="top" align="left">Sub-equatorial</td>
<td valign="top" align="left"><bold>Bacteroidota</bold>: <italic>Myroides</italic> <bold>Proteobacteria</bold><italic>: Plesiomonas, Yersinia, Enterobacter, Shewanella, Aeromonas, Vibrio</italic></td>
<td valign="top" align="left">Wu et al., <xref ref-type="bibr" rid="B92">2010</xref></td>
</tr> <tr style="background-color:#dee1e1">
<td valign="top" align="left" colspan="6"><bold>Cyprinids</bold></td>
</tr> <tr>
<td valign="top" align="left">Bigheaded carps (<italic>Hypophthalmichthys</italic> spp.)</td>
<td valign="top" align="left">Wild caught</td>
<td valign="top" align="left">Transdanubian, Hungary</td>
<td valign="top" align="left">Temperate</td>
<td valign="top" align="left"><bold>Fusobacteriota</bold><italic>: Cetobacterium</italic>, <bold>Proteobacteria</bold><italic>: Pelomonas, Herbaspirillum, Aeromonas, Shewanella</italic></td>
<td valign="top" align="left">Borsodi et al., <xref ref-type="bibr" rid="B10">2017</xref></td>
</tr> <tr>
<td valign="top" align="left">Crucian carp (<italic>Carassius auratus</italic>)</td>
<td valign="top" align="left">Wild caught</td>
<td valign="top" align="left">Yangtze River basin, China</td>
<td valign="top" align="left">Temperate</td>
<td valign="top" align="left"><bold>Firmicutes</bold><italic>: Clostridium</italic> XI, <bold>Fusobacteriota</bold><italic>: Cetobacterium, Fusobacterium</italic>, <bold>Proteobacteria</bold><italic>: Aeromonas, Chitinibacter, Pseudomonas, Vibrio, Serratia</italic></td>
<td valign="top" align="left">Li et al., <xref ref-type="bibr" rid="B46">2023</xref></td>
</tr> <tr>
<td valign="top" align="left">Bighead carp (<italic>Hypophthalmichthys nobilis</italic>), Silver carp (<italic>Hypophthalmichthys molitrix</italic>), Common carp (<italic>Cyprinus carpio</italic>), Goldfish (<italic>Carassius auratus</italic>), Freshwater drum (<italic>Aplodinotus grunniens</italic>)</td>
<td valign="top" align="left">Farm-raised and wild caught</td>
<td valign="top" align="left">Illinois River, USA</td>
<td valign="top" align="left">Temperate</td>
<td valign="top" align="left"><bold>Firmicutes</bold>: Abundant <bold>Fusobacteriota</bold>: Most Abundant<sup>&#x0002A;</sup> <bold>Proteobacteria</bold>: Abundant</td>
<td valign="top" align="left">Eichmiller et al., <xref ref-type="bibr" rid="B22">2016</xref></td>
</tr> <tr>
<td valign="top" align="left">Silver carp (<italic>Hypophthalmichthys molitrix</italic>)</td>
<td valign="top" align="left">Wild caught</td>
<td valign="top" align="left">Havana, IL, Louisiana, MO, West Lafayette, IN, and McBaine, MO, USA</td>
<td valign="top" align="left">Temperate</td>
<td valign="top" align="left"><bold>Firmicutes</bold><italic>: Bacillus, Clostridium</italic>, <bold>Proteobacteria</bold><italic>: Aeromonas, Enterobacter</italic></td>
<td valign="top" align="left">Ye et al., <xref ref-type="bibr" rid="B97">2014</xref></td>
</tr> <tr>
<td valign="top" align="left">Bighead carp (<italic>Hypophthalmichthys nobilis</italic>), Silver carp (<italic>Hypophthalmichthys molitrix</italic>), common carp (<italic>Cyprinus carpio</italic>)</td>
<td valign="top" align="left">Farm-raised and wild caught</td>
<td valign="top" align="left">Illinois River, USA</td>
<td valign="top" align="left">Sub-tropical</td>
<td valign="top" align="left"><bold>Firmicutes</bold>: Abundant, <bold>Fusobacteriota</bold>: Most Abundant,<sup>&#x0002A;</sup> <bold>Proteobacteria</bold>: Abundant</td>
<td valign="top" align="left">Eichmiller et al., <xref ref-type="bibr" rid="B22">2016</xref></td>
</tr> <tr>
<td valign="top" align="left">Crucian carp (<italic>Carassius auratus</italic>)</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">Jiangsu, China</td>
<td valign="top" align="left">Sub-tropical</td>
<td valign="top" align="left"><bold>Firmicutes</bold><italic>: Holdemania, Lactococcus, Staphylococcus</italic>, <bold>Fusobacteriota</bold>: Cetobacterium, <bold>Proteobacteria</bold><italic>: Vibrio, Aeromonas, Shewanella</italic></td>
<td valign="top" align="left">Li et al., <xref ref-type="bibr" rid="B43">2017a</xref></td>
</tr> <tr>
<td valign="top" align="left">Herbivorous grass carp (<italic>Ctenopharyngodon Idellus</italic>) and <italic>Carnivorous Siniperca chuatsi</italic>, and <italic>Silurus meridionalis</italic>)</td>
<td valign="top" align="left">Wild caught</td>
<td valign="top" align="left">Wuhan, China</td>
<td valign="top" align="left">Sub-tropical</td>
<td valign="top" align="left"><bold>Bacteroidota</bold><italic>: Bacteroides</italic>, <bold>Firmicutes</bold><italic>: Lactococcus, Clostridium, Proteocatella, Anaerorhabdus, Clostridium</italic>, <bold>Proteobacteria</bold>: <italic>Acinetobacte, Aeromonas, Serratia, Steroidobacter, Dechloromonas</italic></td>
<td valign="top" align="left">Yan et al., <xref ref-type="bibr" rid="B96">2016</xref></td>
</tr> <tr>
<td valign="top" align="left">Cyprinid Fishes herbivorous grass carp (<italic>Ctenopharyngodon idellus</italic>) and blunt snout bream (<italic>Megalobrama amblycephala</italic>); omnivorous crucian carp (<italic>Carassius auratus</italic>); filter:feeding silver carp (<italic>Hypophthalmichthys molitrix</italic>) and bighead carp (<italic>Hypophthalmichthys nobilis</italic>)</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">Wuhan, China</td>
<td valign="top" align="left">Sub-tropical</td>
<td valign="top" align="left"><bold>Proteobacteria:</bold> <italic>Vibrio, Aeromonas, Shewanella</italic></td>
<td valign="top" align="left">Li et al., <xref ref-type="bibr" rid="B45">2017b</xref></td>
</tr> <tr>
<td valign="top" align="left">Transgenic common carp (<italic>Cyprinus carpio</italic> L.)</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">Guanqiao, China</td>
<td valign="top" align="left">Sub-tropical</td>
<td valign="top" align="left"><bold>Bacteroidota</bold>: Abundant <bold>Firmicutes</bold>: Abundant</td>
<td valign="top" align="left">Li et al., <xref ref-type="bibr" rid="B47">2013</xref></td>
</tr> <tr>
<td valign="top" align="left">Grass carp (<italic>Ctenopharyngodon idellus</italic>)</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">Jingzhou, Hubei, China</td>
<td valign="top" align="left">Sub-tropical</td>
<td valign="top" align="left"><bold>Actinobacteriota</bold><italic>: Actinomyces</italic> <bold>Firmicutes</bold><italic>: Clostridium</italic> <bold>Proteobacteria</bold><italic>: Citrobacter</italic></td>
<td valign="top" align="left">Wu et al., <xref ref-type="bibr" rid="B93">2012</xref></td>
</tr> <tr>
<td valign="top" align="left">Grass carp (<italic>Ctenopharyngodon idellus</italic>), Crucian carp (<italic>Carassius cuvieri</italic>), and Bighead carp (<italic>Hypophthalmichthys nobilis</italic>)</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">Wuhan, China</td>
<td valign="top" align="left">Sub-tropical</td>
<td valign="top" align="left"><bold>Bacteroidota</bold><italic>: Bacteroides</italic>, <bold>Firmicutes</bold><italic>: Clostridium, Proteocatella</italic> <bold>Fusobacteriota</bold><italic>: Cetobacterium</italic>, <bold>Proteobacteria</bold><italic>: Aeromonas</italic></td>
<td valign="top" align="left">Li et al., <xref ref-type="bibr" rid="B44">2015</xref></td>
</tr> <tr>
<td valign="top" align="left">Gibel carp (<italic>Carassius auratus gibelio</italic>)</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">Wuhan, China</td>
<td valign="top" align="left">Sub-tropical</td>
<td valign="top" align="left"><bold>Actinobacteriota</bold>: <italic>Catellibacterium</italic>, <bold>Firmicutes</bold>: <italic>Cetobacterium, Holdemania, Lactococcus, Staphylococcus</italic> <bold>Proteobacteria</bold>: <italic>Pseudomonas, Acinetobacter, Serratia, Shewanella, Aeromonas, Roseomonas, Ensifer, Bose</italic></td>
<td valign="top" align="left">Li et al., <xref ref-type="bibr" rid="B48">2017c</xref></td>
</tr> <tr>
<td valign="top" align="left">Grass carp</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">Wuhan, China</td>
<td valign="top" align="left">Sub-equatorial</td>
<td valign="top" align="left"><bold>Actinobacteriota</bold><italic>: Adlercreutzia</italic>, <bold>Bacteroidota</bold><italic>: Chryseobacterium, Citrobacter</italic>, <bold>Firmicutes</bold><italic>: Enterococcus</italic></td>
<td valign="top" align="left">Xiong et al., <xref ref-type="bibr" rid="B94">2022</xref></td>
</tr> <tr>
<td valign="top" align="left">Grass carp (<italic>Ctenopharyngodon Idellus</italic>)</td>
<td valign="top" align="left">Wild caught</td>
<td valign="top" align="left">Wuhan, China</td>
<td valign="top" align="left">Sub-equatorial</td>
<td valign="top" align="left"><bold>Firmicutes</bold><italic><bold>:</bold> Lactococcus, Leuconostoc, Weisella</italic></td>
<td valign="top" align="left">Yan et al., <xref ref-type="bibr" rid="B96">2016</xref></td>
</tr> <tr>
<td valign="top" align="left"><italic>M. amblycephala</italic> and <italic>C. idellus, S. chuatsi</italic> and <italic>C. alburnus</italic>, omnivorous C. <italic>carpio and C. auratus</italic></td>
<td valign="top" align="left">Wild caught</td>
<td valign="top" align="left">Wuhan, China</td>
<td valign="top" align="left">Sub-equatorial</td>
<td valign="top" align="left"><bold>Firmicutes</bold><italic>: Clostridium</italic> <bold>Fusobacteriota</bold>: <italic>Leptotrichia, Cetobacterium</italic> <bold>Proteobacteria</bold>: <italic>Citrobacter</italic></td>
<td valign="top" align="left">Liu et al., <xref ref-type="bibr" rid="B49">2016</xref></td>
</tr> <tr>
<td valign="top" align="left">Transgenic Common Carp (<italic>Cyprinus carpio L</italic>.)</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">Guanqiao, China</td>
<td valign="top" align="left">Sub-equatorial</td>
<td valign="top" align="left"><bold>Bacteroidota</bold>: Abundant <bold>Firmicutes</bold>: Most Abundant<sup>&#x0002A;</sup></td>
<td valign="top" align="left">Li et al., <xref ref-type="bibr" rid="B47">2013</xref></td>
</tr> <tr>
<td valign="top" align="left">Indian major carps (IMCs), rohu (<italic>Labeo rohita</italic>), catla (<italic>Catla catla</italic>) and mrigal (<italic>Cirrhinus mrigala</italic>)</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">West Bengal, India</td>
<td valign="top" align="left">Tropical</td>
<td valign="top" align="left"><bold>Fusobacteriota</bold>: <italic>Fusobacterium</italic>, <bold>Proteobacteria</bold>: <italic>Aeromonas</italic></td>
<td valign="top" align="left">Mukherjee et al., <xref ref-type="bibr" rid="B63">2020</xref></td>
</tr> <tr>
<td valign="top" align="left">Catla <italic>(Cyprinus catla)</italic>, Common carp (<italic>Cyprinus carpio)</italic>, mrigal <italic>(Cyprinus mrigala)</italic> and Rohu <italic>(Labeo rohita)</italic></td>
<td valign="top" align="left">Wild caught</td>
<td valign="top" align="left">Maharashtra, India</td>
<td valign="top" align="left">Tropical</td>
<td valign="top" align="left"><bold>Actinobacteriota</bold>: Abundant <bold>Firmicutes</bold>: <italic>Bacillus, Clostridium, Lactococcus</italic> <bold>Proteobacteria</bold>: <italic>Sphingomonas</italic></td>
<td valign="top" align="left">Pingle and Khandagle, <xref ref-type="bibr" rid="B66">2023</xref></td>
</tr> <tr style="background-color:#dee1e1">
<td valign="top" align="left" colspan="6"><bold>Cichlidae (Tilapia)</bold></td>
</tr> <tr>
<td valign="top" align="left"><italic>Amphilophus</italic> sp.</td>
<td valign="top" align="left">Wild caught</td>
<td valign="top" align="left">Nicaragua and Maganua, USA</td>
<td valign="top" align="left">Tropical</td>
<td valign="top" align="left"><bold>Proteobacteria</bold>: Most Abundant<sup>&#x0002A;</sup> <bold>Fusobacteria</bold>: Most Abundant<sup>&#x0002A;</sup> <bold>Firmicutes</bold>: Abundant <bold>Bacteroidetes</bold>: Abundant <bold>Planctomycetes</bold>: Abundant</td>
<td valign="top" align="left">Baldo et al., <xref ref-type="bibr" rid="B6">2019</xref></td>
</tr> <tr>
<td valign="top" align="left">African cichlid</td>
<td valign="top" align="left">Wild caught</td>
<td valign="top" align="left">Tanganyika, Zambia; Barombi Mbo, Cameroon</td>
<td valign="top" align="left">Sub-equatorial</td>
<td valign="top" align="left"><bold>Fusobacteria</bold>: <italic>Plecodus</italic> sp.<italic>, Xenotilapia</italic> sp.<italic>, Myaka</italic> sp. <bold>Proteobacteria</bold>: <italic>Lamprologus</italic> sp.<italic>, Variabilichromis</italic> sp.<italic>, Sarotherodon</italic> sp., <bold>Firmicutes</bold>: <italic>Lepidiolamprologus</italic> sp.<italic>, Neolamprologus</italic> sp.<italic>, Sarotherodon</italic> sp. <bold>Planctomycetes</bold>: <italic>Lepidiolamprologus</italic> sp.<italic>s, Interochromis</italic> sp.<italic>, Konia</italic> sp. <bold>Actinobacteria</bold>: <italic>Altolamprologus</italic> sp.<italic>, Ophthalmotilapia</italic> sp.<italic>, Sarotherodon</italic> sp. <bold>Verrucomicrobia</bold>: <italic>Enantiopus</italic> sp.<italic>, Eretmodus</italic> sp. <bold>Chlamydiae</bold>: <italic>Gnathochromis</italic> sp.<italic>, Simochromis</italic> sp. <bold>Bacteroidetes</bold>: <italic>Cyprichromis</italic> sp.<italic>, Konia</italic> sp. <bold>Chloroflexi</bold>: <italic>Neolamprologus</italic> sp.<italic>, Pungu</italic> sp.</td>
<td valign="top" align="left">Baldo et al., <xref ref-type="bibr" rid="B5">2017</xref></td>
</tr> <tr>
<td/>
<td valign="top" align="left">Wild caught</td>
<td valign="top" align="left">Tanganyika, Zambia and Tanzania</td>
<td valign="top" align="left">Sub-equatorial</td>
<td valign="top" align="left"><bold>Fusobacteria</bold>: <italic>Cetobacterium</italic> <bold>Firmicutes</bold>: <italic>Clostridium, Turicibacter, Clostridium</italic> XI, Lachnospiraceae (family), Clostridiales (order), Clostridiaceae (family), <italic>Bacillus</italic>, <bold>Proteobacteria</bold>: <italic>Plesiomonas, Aeromonas</italic>, Neisseriaceae (family), <italic>Achromobacter</italic>, <bold>Planctomycetes</bold>: Pirellulaceae (family)</td>
<td valign="top" align="left">Baldo et al., <xref ref-type="bibr" rid="B7">2015</xref></td>
</tr> <tr>
<td valign="top" align="left">Nile tilapia (<italic>Oreochromis niloticus</italic>)</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">Darmstadt, Germany</td>
<td valign="top" align="left">Sub-tropical</td>
<td valign="top" align="left"><bold>Actinobacteriota</bold>: <italic>Arthrobacter, Chitinilyticum, Leucobacter, Luteitalea</italic> <bold>Proteobacteria</bold>: Most Abundant<sup>&#x0002A;</sup><italic>Acinetobacter, Aeromonas, Aquabacterium, Dechloromonas, Pseudomonas, Psychrobacter, Reyranella, Shewanella, Stenotrophomonas</italic> <bold>Firmicutes</bold>: <italic>Lactobacillus, Lactococcus, Staphylococcus</italic> <bold>Bacteroidetes</bold>: <italic>Chryseobacterium, Dinghuibacter, Flavobacterium</italic> <bold>Planctomycetes</bold>: <italic>Blastopirellula</italic> <bold>Verrucomicrobia</bold>: Abundant</td>
<td valign="top" align="left">Guimar&#x000E3;es et al., <xref ref-type="bibr" rid="B28">2021</xref></td>
</tr> <tr>
<td valign="top" align="left">Nile tilapia (<italic>Oreochromis niloticus</italic>)</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">Gaozhou, China</td>
<td valign="top" align="left">Sub-tropical</td>
<td valign="top" align="left"><bold>Firmicutes</bold>: <italic>Clostridium_sensu_stricto_2, Faecalibacterium, Hathewaya, f__Clostridiaceae_1_Unclassified, Terrisporobacter</italic> <bold>Proteobacteria</bold>: <italic>Escherichia&#x02013;Shigella, Acinetobacter, Aeromonas</italic> <bold>Bacteroidota</bold>: <italic>f__Muribaculaceae_Unclassified, Bacteroides</italic></td>
<td valign="top" align="left">Kuebutornye et al., <xref ref-type="bibr" rid="B38">2020</xref></td>
</tr> <tr>
<td valign="top" align="left">Nile tilapia (<italic>Oreochromis niloticus</italic>)</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">Charoen, Thailand</td>
<td valign="top" align="left">Tropical</td>
<td valign="top" align="left"><bold>Fusobacteria</bold>: <italic>Cetobacterium</italic> <bold>Proteobacteria</bold>: <italic>Aquaspirillum, Edwardsiella, Plesiomonas, Balneimonas, Rhodobacter</italic> <bold>Firmicutes</bold>: <italic>Weissella, Bacillus, Staphylococcus</italic> <bold>Actinobacteria</bold>: <italic>Corynebacterium</italic></td>
<td valign="top" align="left">Adeoye et al., <xref ref-type="bibr" rid="B1">2016</xref></td>
</tr> <tr>
<td valign="top" align="left">Nile tilapia (<italic>Oreochromis niloticus</italic>)</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">Sonora, Mexico</td>
<td valign="top" align="left">Sub-tropical</td>
<td valign="top" align="left"><bold>Proteobacteria</bold>: Most Abundant<sup>&#x0002A;</sup> <bold>Fusobacteria</bold>: Most Abundant<sup>&#x0002A;</sup> <bold>Actinobacteria</bold>: Abundant <bold>Firmicutes</bold>: Abundant</td>
<td valign="top" align="left">Martinez-Porchas et al., <xref ref-type="bibr" rid="B58">2023</xref></td>
</tr>
<tr>
<td valign="top" align="left">Nile tilapia, (<italic>Oreochromis niloticus</italic>)</td>
<td valign="top" align="left">Farm-raised</td>
<td valign="top" align="left">Plymouth University, UK</td>
<td valign="top" align="left">Temperate</td>
<td valign="top" align="left"><bold>Actinobacteria</bold>: <italic>Mycobacterium, Propionibacterium, Curtobacterium, Phyciococcus, Corynebacterium</italic> <bold>Proteobacteria</bold>: <italic>Acinetobacter, Cobetia, Legionella, Plesiomonas, Janthinobacterium, Sphingomonas, Paracoccus, Methylobacterium, Rhodoplanes, Hyphomicrobium, Bradyrhizobium, Afipia</italic> <bold>Fusobacteria</bold>: <italic>Cetobacterium</italic> <bold>Firmicutes</bold>: <italic>Streptococcus, Weissella, Leuconostoc, Pediococcus, Lactobacillus, Enterococcus, Staphylococcus, Bacillus, Veillonella</italic> <bold>Bacteroidetes</bold>: Other (Order Clostridiales), Other (Family Peptostreptococcaceae) <bold>Spirochaetes</bold>: SMB53 <bold>Cyanobacteria</bold>: <italic>Streptophyta</italic></td>
<td valign="top" align="left">Standen et al., <xref ref-type="bibr" rid="B80">2015</xref></td>
</tr></tbody>
</table>
</table-wrap>
<p>Temperature is another key factor influencing gut microbiome composition across climate zones. Fish from warmer environments often display greater microbial diversity, with temperature playing a crucial role in shaping species-specific responses (Wong and Rawls, <xref ref-type="bibr" rid="B91">2012</xref>; Kokou et al., <xref ref-type="bibr" rid="B36">2018</xref>). For example, yellow-tail kingfish showed higher gut microbiota richness at 26&#x000B0;C than at 20&#x000B0;C (Soriano et al., <xref ref-type="bibr" rid="B78">2018</xref>), while turbot exhibited greater diversity at 20&#x000B0;C (Guerreiro et al., <xref ref-type="bibr" rid="B27">2016</xref>). In rainbow trout, higher temperatures were associated with a reduction in Firmicutes (Huyben et al., <xref ref-type="bibr" rid="B32">2018</xref>), and in salmon, higher temperatures led to a decrease in Acinetobacter and an increase in pathogenic <italic>Vibrio</italic> (Ley et al., <xref ref-type="bibr" rid="B41">2008</xref>). Such variations underscore important role of temperature in influencing microbiota, particularly in temperature-sensitive fish (Chevalier et al., <xref ref-type="bibr" rid="B12">2015</xref>). Among those climatic zones, fish from the subtropical region displayed the highest microbial diversity, with tropical, temperate, and sub-equatorial regions following diversity levels (<xref ref-type="fig" rid="F1">Figures 1D</xref>, <xref ref-type="fig" rid="F1">E</xref>, <xref ref-type="table" rid="T1">Table 1</xref>).</p></sec>
<sec id="s3">
<title>Gut microbiome: an indicator for fish conservation and management strategies</title>
<p>The gut microbiome is increasingly recognized as an important indicator of environmental health and the adaptability of fish populations, offering valuable insights for conservation efforts (Soh et al., <xref ref-type="bibr" rid="B77">2024</xref>). Factors such as geographic location, exposure to contaminants, urbanization, and the introduction of invasive species can significantly disrupt the gut microbiome, impacting essential processes like digestion, metabolism, immunity, and overall health (Zhu et al., <xref ref-type="bibr" rid="B104">2021a</xref>; Clough et al., <xref ref-type="bibr" rid="B13">2023</xref>; Lennon et al., <xref ref-type="bibr" rid="B40">2023</xref>; Lorgen-Ritchie et al., <xref ref-type="bibr" rid="B52">2023</xref>). Such disruptions complicate conservation strategies, particularly for endangered species, by altering microbial diversity and reducing adaptability. Shifts in microbial diversity in response to environmental pollutants or habitat changes can provide early warnings of ecological imbalance, facilitating timely conservation interventions (Zhu et al., <xref ref-type="bibr" rid="B104">2021a</xref>).</p>
<p>Invasive species, such as Nile tilapia, illustrate how microbiome diversity can confer competitive advantages. Compared to native fish, invasive tilapia display higher gut microbial alpha diversity, reduced interspecies microbial competition, and enhanced food utilization, supporting niche expansion and local adaptation (Gu et al., <xref ref-type="bibr" rid="B26">2020</xref>). Similarly, bighead carp and silver carp, major aquaculture species in East Asia, have become invasive in North America, where hybridization in the Mississippi River Basin (MRB) has further increased their adaptability (Wang et al., <xref ref-type="bibr" rid="B89">2020</xref>). Studies suggest that hybrids benefit from a diversified gut microbiome, which, along with genomic adaptability, may facilitate invasion by supporting survival and local adaptation (Wang et al., <xref ref-type="bibr" rid="B89">2020</xref>; Zhu et al., <xref ref-type="bibr" rid="B105">2021b</xref>). These findings underscore the critical role of the gut microbiome in driving ecological success and adaptability in invasive species. As environmental pressures intensify, understanding the interactions between host genetics and microbiome diversity will be essential for managing invasive populations and protecting native biodiversity.</p>
<p>Managing gut microbiota in captive breeding programs can improve reintroduction success rates by enhancing the resilience and ecological fitness of released fish in their natural habitats (Zhu et al., <xref ref-type="bibr" rid="B104">2021a</xref>). Microbiome-based interventions, including probiotics and prebiotics, hold considerable promise for enhancing resilience and adaptability in both farmed and wild fish, reducing mortality rates and reinforcing conservation outcomes (Jin Song et al., <xref ref-type="bibr" rid="B33">2019</xref>; Vargas-albores et al., <xref ref-type="bibr" rid="B87">2021</xref>; de Jonge et al., <xref ref-type="bibr" rid="B16">2022</xref>). This approach is essential for species recovery, as a well-balanced gut microbiome strengthens adaptability, immune function, and overall health in reintroduced populations, enabling them to thrive in challenging environments.</p>
<p>Advancing sustainable aquaculture and implementing effective conservation strategies for vulnerable fish populations hinges on a comprehensive understanding of gut microbiome dynamics. Research has shown that gut microbiota plays a fundamental role in host resilience, immune function, and adaptation to environmental changes, all of which are critical for both farmed and wild fish (Fonseca and Fuentes, <xref ref-type="bibr" rid="B23">2023</xref>; Zhu and Wang, <xref ref-type="bibr" rid="B103">2023</xref>). Fish migration and breeding activities are closely associated with variations in gut microbiome composition, as they drive physiological and environmental changes (Llewellyn et al., <xref ref-type="bibr" rid="B51">2016</xref>; Hamilton et al., <xref ref-type="bibr" rid="B29">2019</xref>; Liu et al., <xref ref-type="bibr" rid="B50">2021</xref>). As conservation and aquaculture practices increasingly incorporate microbiome management, this could become a powerful tool for sustaining biodiversity, aiding species recovery, and supporting sustainable aquaculture.</p>
<p>Embracing microbiome-based solutions strengthens the health and adaptability of individual species and contributes to the stability of entire aquatic ecosystems. Such microbiome-focused approaches have the potential to transform conservation practices by enhancing species survival, mitigating the impacts of invasive species, and restoring ecosystem health. By integrating these strategies, conservation efforts can foster resilient and balanced aquatic environments, paving the way for the long-term sustainability of our aquatic ecosystems.</p></sec>
<sec id="s4">
<title>Discussion and future directions</title>
<p>This review provides a global perspective on the gut microbiome of four major aquaculture fish groups: cyprinids, ictalurids (catfish), cichlids (tilapia), and salmonids. It highlighted microbial composition and diversity across geographic regions and contrasting farmed and wild environmental conditions (<xref ref-type="fig" rid="F1">Figure 1</xref>). By focusing on these important species, we identified dominant phyla and critical patterns that consistently appear across studies. Notably, samples from temperate, sub-equatorial, and subtropical zones exhibited the highest microbial diversity, emphasizing the interplay between taxonomic and environmental factors in shaping the microbiome (<xref ref-type="fig" rid="F1">Figure 1E</xref>, <xref ref-type="table" rid="T1">Table 1</xref>). These findings underscore the adaptive significance of the gut microbiome in supporting essential functions such as digestion, immunity, and overall health, thereby enhancing the resilience and productivity of aquaculture species.</p>
<sec>
<title>Limitations and opportunities</title>
<p>While this review provides valuable insights, it is important to acknowledge its limitations. The selection of species and groups, while reasonable, does not fully represent the diversity of fish (Riera and Baldo, <xref ref-type="bibr" rid="B71">2020</xref>). This limitation may restrict the generalizability of our findings, as different species and habitats harbor distinct microbiome compositions. Even within a single taxonomic group, various species inhabiting different niches can possess diverse gut microbiomes, which might have been underestimated in our review. Future research should aim to expand the analysis to include all species and taxonomic groups studied.</p>
<p>Additionally, the focus on microbial composition at the phylum level, while informative, represents a much higher taxonomic resolution. Comparisons at the genus or even at the species level would offer more detailed and biologically meaningful insights, particularly for understanding functional relationships within the microbiome. Furthermore, variability in sampling methods, sequencing depth, and environmental contexts across the reviewed studies introduces potential biases. This heterogeneity complicates cross-study comparisons and limits the ability to draw broad, definitive conclusions. A meta-analysis of raw sequencing data could address these inconsistencies, yielding more robust and statistically validated insights.</p>
<p>A geographic bias in sampling is evident, with most studies conducted in North America and East Asia, while regions such as Central Europe and Africa are underrepresented (<xref ref-type="fig" rid="F1">Figure 1D</xref>, <xref ref-type="table" rid="T1">Table 1</xref>). This imbalance, similar to patterns observed in the Earth Microbiome Project (Gilbert et al., <xref ref-type="bibr" rid="B25">2018</xref>), underscores the need for increased geographic diversity to achieve a truly global perspective on fish gut microbiomes. Expanding research efforts to include diverse regions and underrepresented species will be crucial for creating a comprehensive and unbiased framework for understanding and leveraging microbiome data in aquaculture and conservation.</p></sec>
<sec>
<title>Addressing microbiome complexities with innovative, integrated approaches</title>
<p>The study of fish gut microbiomes is inherently complex due to the various influencing variables, such as diet, water quality, temperature, and salinity, which complicate efforts to isolate specific factors affecting microbial community structure (Egerton et al., <xref ref-type="bibr" rid="B21">2018</xref>). The gut microbiome interacts dynamically with the host and the environment, requiring research to move beyond simple associations to uncover complex causal relationships (Xiong et al., <xref ref-type="bibr" rid="B95">2019</xref>). The functional impact of the microbiome depends on the entire ecological network, where diet, environmental factors, and microbial composition interact in interdependent ways (Talwar et al., <xref ref-type="bibr" rid="B83">2018</xref>; Diwan et al., <xref ref-type="bibr" rid="B19">2022</xref>).</p>
<p>To address these challenges, the holobiont approach and multi-omics techniques should be employed. Future research on fish can follow the lead of large-scale microbiome initiatives, such as the Human Microbiome Project (Turnbaugh et al., <xref ref-type="bibr" rid="B86">2007</xref>), to advance fish microbiome studies. For instance, the holobiont model has been used to explore how microbiomes interact with host genomes to drive adaptability and invasiveness in hybrid bighead and silver carp within the Mississippi River Basin (Wang et al., <xref ref-type="bibr" rid="B89">2020</xref>; Zhu et al., <xref ref-type="bibr" rid="B105">2021b</xref>). The integration of multi-omics data can uncover the functional roles of microbiomes in fish health and adaptation by linking microbial genes and metabolic pathways to host physiological traits, such as digestion efficiency, immunity, and stress tolerance.</p>
<p>Beyond experimental and field research, computational modeling and theoretical studies are crucial for enhancing our understanding of microbiomes as integral components of complex ecological networks (Kumar et al., <xref ref-type="bibr" rid="B39">2019</xref>). Modeling microbial interaction networks, predicting the responses of these networks to environmental changes, and simulating diverse scenarios to assess the far-reaching impacts of the microbiome on host resilience and aquaculture productivity are promising avenues for future work. By combining experimental and computational approaches, researchers can unravel the intricate interdependencies among microbiomes, host, and environments, providing deeper insights into ecosystem functioning and advancing practical applications.</p></sec>
<sec>
<title>Perspectives in aquaculture and conservation</title>
<p>Aquaculture practices, such as integrated pond fish farming and natural fish germplasm resource conservation, have been pivotal in sustainable aquaculture in regions like China (Li et al., <xref ref-type="bibr" rid="B42">1990</xref>; Lu et al., <xref ref-type="bibr" rid="B53">1997</xref>, <xref ref-type="bibr" rid="B55">2020</xref>). Microbiome is likely to play a crucial role in these processes. Comparative analyses of gut microbial communities between natural and pond-cultured populations, as well as within the same pond ecosystem, could unveil the underlying mechanisms.</p>
<p>Further studies on the role of the microbiome in fish nutrition and health are necessary to enhance aquaculture productivity and promote healthier aquatic ecosystems. Developing microbiome-targeted feeds enriched with prebiotics, probiotics, or synbiotics can stimulate beneficial gut microbes and improve fish growth and disease resistance. For example, utilizing probiotics and incorporating alternative protein sources, such as insect meals, can enhance gut health and optimize aquaculture practices (Fonseca and Fuentes, <xref ref-type="bibr" rid="B23">2023</xref>; Hasan et al., <xref ref-type="bibr" rid="B30">2023</xref>). Feed formulations should be optimized based on microbiome profiles to improve nutrient uptake and resilience to pathogens. Insights from studies on wild and hybrid species can inform the design of functional feeds and strategies to enhance aquaculture productivity (Reshma et al., <xref ref-type="bibr" rid="B70">2018</xref>; Cui et al., <xref ref-type="bibr" rid="B15">2022</xref>).</p>
<p>Applying microbiome research to improve recirculating aquaculture systems by optimizing microbial communities in biofilters and water systems is crucial (Rurangwa and Verdegem, <xref ref-type="bibr" rid="B72">2015</xref>; Mugwanya et al., <xref ref-type="bibr" rid="B62">2021</xref>). Studying sediment and water column microbiomes to enhance nutrient recycling and minimize environmental impact is also essential. Promoting integrated multi-trophic aquaculture systems that leverage microbiome interactions across species can contribute to sustainable aquaculture practices (Troell et al., <xref ref-type="bibr" rid="B85">2009</xref>).</p>
<p>Microbiome research is critical for understanding and maintaining wild fish populations. Future research should study the microbiomes of wild fish populations to understand their role in species health and ecosystem stability. Using microbiome monitoring to support species reintroduction programs and mitigate the impacts of invasive species is crucial. Enhancing biodiversity conservation by protecting critical microbial symbionts associated with endangered species is imperative.</p></sec></sec>
<sec sec-type="conclusions" id="s5">
<title>Conclusion</title>
<p>The fish gut microbiome represents a promising frontier for advancements in aquaculture and conservation biology. By harnessing the power of microbiome research, we can develop more sustainable aquaculture practices, enhance the resilience of wild fish populations, and safeguard the delicate balance of aquatic ecosystems. Interdisciplinary approaches and innovative technologies will pave the way for transformative solutions to global challenges in aquaculture sustainability and biodiversity conservation.</p></sec>
</body>
<back>
<sec sec-type="author-contributions" id="s6">
<title>Author contributions</title>
<p>NK: Writing &#x02013; original draft, Writing &#x02013; review &#x00026; editing. NL: Writing &#x02013; review &#x00026; editing. JK: Writing &#x02013; review &#x00026; editing. HC: Writing &#x02013; review &#x00026; editing. GL: Writing &#x02013; review &#x00026; editing, Conceptualization, Writing &#x02013; original draft. JW: Conceptualization, Writing &#x02013; original draft, Writing &#x02013; review &#x00026; editing. CW: Conceptualization, Writing &#x02013; original draft, Writing &#x02013; review &#x00026; editing.</p>
</sec>
<sec sec-type="funding-information" id="s7">
<title>Funding</title>
<p>The author(s) declare financial support was received for the research, authorship, and/or publication of this article. CW is grateful for funding support from the National Natural Science Foundation of China under grant number: 32172959. GL acknowledges funding support from the Mississippi River Basin Panel on Aquatic Nuisance Species for the Invasive Carp Genetics Project.</p>
</sec>
<ack><p>This review is dedicated in part to the late Louise Bernatchez, our esteemed mentor and long-time collaborator, and Sifa Li for his pioneering work in conserving the four major Chinese carp species. We extend our gratitude to L. Zhu for his foundational contributions and for inviting us to undertake this review. We also thank our lab members for their insightful comments and invaluable contributions.</p>
</ack>
<sec sec-type="COI-statement" id="conf1">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec sec-type="ai-statement" id="s8">
<title>Generative AI statement</title>
<p>The author(s) declare that Gen AI was used in the creation of this manuscript. We used generative AI tools to help with English and improve language.</p></sec>
<sec sec-type="disclaimer" id="s9">
<title>Publisher&#x00027;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
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