Abstract
Intestinal flora is closely related to the health of organisms and the occurrence and development of diseases. The study of intestinal flora will provide a reference for the research and treatment of disease pathogenesis. Upon hatching, fish begin to acquire a microbial community in the intestine. In response to the environment and the host itself, the fish gut eventually develops a unique set of microflora, with some microorganisms being common to different fish. The existence of intestinal microorganisms creates an excellent microecological environment for the host, while the fish symbiotically provides conditions for the growth and reproduction of intestinal microflora. The intestinal flora and the host are interdependent and mutually restrictive. This review mainly describes the formation of fish intestinal flora, the function of normal intestinal flora, factors affecting intestinal flora, and a series of fish models.
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References
Arias-Jayo N, Abecia L, Alonso-Sáez L, Ramirez-Garcia A, Rodriguez A, Pardo MA (2018) High-fat diet consumption induces microbiota dysbiosis and intestinal inflammation in zebrafish. Microb Ecol 76:1089–1101. https://doi.org/10.1007/s00248-018-1198-9
Bakke I, Coward E, Andersen T, Vadstein O (2015) Selection in the host structures the microbiota associated with developing cod larvae (Gadus morhua). Environ Microbiol 17:3914–3924. https://doi.org/10.1111/1462-2920.12888
Baldo L, Riera JL, Salzburger W, Barluenga M (2019) Phylogeography and ecological niche shape the cichlid fish gut microbiota in Central American and African Lakes Frontiers in microbiology 10:2372 https://doi.org/10.3389/fmicb.2019.02372
Baldo L, Riera JL, Tooming-Klunderud A, Albà MM, Salzburger W (2015) Gut Microbiota Dynamics during Dietary Shift in Eastern African Cichlid Fishes. PloS One 10:e0127462. https://doi.org/10.1371/journal.pone.0127462
Bates JM, Mittge E, Kuhlman J, Baden KN, Cheesman SE, Guillemin K (2006) Distinct signals from the microbiota promote different aspects of zebrafish gut differentiation. Dev Biol 297:374–386. https://doi.org/10.1016/j.ydbio.2006.05.006
Bledsoe JW, Peterson BC, Swanson KS, Small BC (2016) Ontogenetic characterization of the intestinal microbiota of channel catfish through 16S rRNA gene sequencing reveals insights on temporal shifts and the influence of environmental microbes. PloS one 11:e0166379. https://doi.org/10.1371/journal.pone.0166379
Boulangé CL, Neves AL, Chilloux J, Nicholson JK, Dumas ME (2016) Impact of the gut microbiota on inflammation, obesity, and metabolic disease. Genome Med 8:42. https://doi.org/10.1186/s13073-016-0303-2
Brugman S et al (2014) T lymphocytes control microbial composition by regulating the abundance of Vibrio in the zebrafish gut. Gut Microbes 5:737–747. https://doi.org/10.4161/19490976.2014.972228
Burtseva O, Baulina O, Zaytseva A, Fedorenko T, Chekanov K, Lobakova E (2020) In vitro biofilm formation by bioluminescent bacteria isolated from the marine fish gut. Microb Ecol. https://doi.org/10.1007/s00248-020-01652-0
Catalán N, Villasante A, Wacyk J, Ramírez C, Romero J (2018) Fermented soybean meal increases lactic acid bacteria in gut microbiota of Atlantic salmon (Salmo salar). Probiotics Antimicrob Proteins 10:566–576. https://doi.org/10.1007/s12602-017-9366-7
Chapagain P, Arivett B, Cleveland BM, Walker DM, Salem M (2019) Analysis of the fecal microbiota of fast- and slow-growing rainbow trout (Oncorhynchus mykiss). BMC Genomics 20:788. https://doi.org/10.1186/s12864-019-6175-2
Daly K et al (2019) Host selectively contributes to shaping intestinal microbiota of carnivorous and omnivorous fish. J General App Microbiol 65:129–136. https://doi.org/10.2323/jgam.2018.07.003
Deb S, Das L, Das SK (2020) Composition and functional characterization of the gut microbiome of freshwater pufferfish (Tetraodon cutcutia). Arch microbiol 202:2761–2770. https://doi.org/10.1007/s00203-020-01997-7
DeBofsky A, Xie Y, Jardine TD, Hill JE, Jones PD, Giesy JP (2020) Effects of the husky oil spill on gut microbiota of native fishes in the North Saskatchewan River. Canada Aquatic Toxicol (amsterdam, Netherlands) 229:105658. https://doi.org/10.1016/j.aquatox.2020.105658
Ding Q et al (2018) The hepatotoxicity of palmitic acid in zebrafish involves the intestinal microbiota. J Nutr 148:1217–1228. https://doi.org/10.1093/jn/nxy084
Dooley K, Zon LI (2000) Zebrafish: a model system for the study of human disease. Curr Opinion Genetics Develop 10:252–256. https://doi.org/10.1016/s0959-437x(00)00074-5
Dubos R, Schaedler RW, Costello R, Hoet P (1965) Indigenous, normal, and autochthonous flora of the gastrointestinal tract. J Exp Med 122:67–76. https://doi.org/10.1084/jem.122.1.67
Faith JJ et al (2013) The long-term stability of the human gut microbiota. Sci (New York, NY) 341:1237439. https://doi.org/10.1126/science.1237439
Falcinelli S et al (2015) Lactobacillus rhamnosus lowers zebrafish lipid content by changing gut microbiota and host transcription of genes involved in lipid metabolism. Sci Rep 5:9336. https://doi.org/10.1038/srep09336
Feng S, Zeng Y, Cai Z, Wu J, Chan LL, Zhu J, Zhou J (2021) Polystyrene microplastics alter the intestinal microbiota function and the hepatic metabolism status in marine medaka (Oryzias melastigma). Sci Total Environ 759:143558. https://doi.org/10.1016/j.scitotenv.2020.143558
Fietz K et al (2018) Mind the Gut: Genomic Insights to Population Divergence and Gut Microbial Composition of Two Marine Keystone Species. Microbiome 6:82. https://doi.org/10.1186/s40168-018-0467-7
Fogarty C, Burgess CM, Cotter PD, Cabrera-Rubio R, Whyte P, Smyth C, Bolton DJ (2019) Diversity and composition of the gut microbiota of Atlantic salmon (Salmo salar) farmed in Irish waters. J App Microbiol 127:648–657. https://doi.org/10.1111/jam.14291
Fu J et al (2017) Aquatic animals promote antibiotic resistance gene dissemination in water via conjugation: role of different regions within the zebra fish intestinal tract, and impact on fish intestinal microbiota. Molecular ecology 26:5318–5333. https://doi.org/10.1111/mec.14255
Gajardo K, Rodiles A, Kortner TM, Krogdahl Å, Bakke AM, Merrifield DL, Sørum H (2016) A high-resolution map of the gut microbiota in Atlantic salmon (Salmo salar): a basis for comparative gut microbial research. Scientific reports 6:30893. https://doi.org/10.1038/srep30893
Galindo-Villegas J, García-Moreno D, de Oliveira S, Meseguer J, Mulero V (2012) Regulation of immunity and disease resistance by commensal microbes and chromatin modifications during zebrafish development. Proc Natl Acad Sci USA 109:E2605-2614. https://doi.org/10.1073/pnas.1209920109
Ganz J (2018) Gut feelings: studying enteric nervous system development, function, and disease in the zebrafish model system Developmental dynamics : an official publication of the American. Association of Anatomists 247:268–278. https://doi.org/10.1002/dvdy.24597
Gaulke CA, Martins ML, Watral VG, Humphreys IR, Spagnoli ST, Kent ML, Sharpton TJ (2019) A longitudinal assessment of host-microbe-parasite interactions resolves the zebrafish gut microbiome's link to pseudocapillaria tomentosa infection and pathology. Microbiome 7:10. https://doi.org/10.1186/s40168-019-0622-9
Giatsis C, Sipkema D, Smidt H, Verreth J, Verdegem M (2014) The colonization dynamics of the gut microbiota in Tilapia Larvae. PloS One 9:e103641. https://doi.org/10.1371/journal.pone.0103641
Goessling W, Sadler KC (2015) Zebrafish: an important tool for liver disease research. Gastroenterol 149:1361–1377. https://doi.org/10.1053/j.gastro.2015.08.034
Gu H, Wang S, Wang X, Yu X, Hu M, Huang W, Wang Y (2020) Nanoplastics impair the intestinal health of the juvenile large yellow croaker Larimichthys crocea. J Hazard Mater 397:122773. https://doi.org/10.1016/j.jhazmat.2020.122773
Guo X, Ran C, Zhang Z, He S, Jin M, Zhou Z (2017) The growth-promoting effect of dietary nucleotides in fish is associated with an intestinal microbiota-mediated reduction in energy expenditure. J Nutr 147:781–788. https://doi.org/10.3945/jn.116.245506
He X, Chaganti SR, Heath DD (2018) Population-specific responses to interspecific competition in the gut microbiota of two Atlantic salmon (Salmo salar) populations. Microbial Ecol 75:140–151. https://doi.org/10.1007/s00248-017-1035-6
Heys C et al. (2020) Neutral processes dominate microbial community assembly in Atlantic salmon, Salmo salar Applied and environmental microbiology 86 https://doi.org/10.1128/aem.02283-19
Huang W et al (2018) Community composition, diversity, and metabolism of intestinal microbiota in cultivated European eel (Anguilla anguilla). App Microbiol Biotechnol 102:4143–4157. https://doi.org/10.1007/s00253-018-8885-9
Jiang M et al (2020) The intestinal microbiota of lake anchovy varies according to sex, body size, and local habitat in Taihu Lake. China MicrobiologyOpen 9:e00955. https://doi.org/10.1002/mbo3.955
Jones J et al (2018) The microbiome of the gastrointestinal tract of a range-shifting marine herbivorous fish. Front Microbiol 9:2000. https://doi.org/10.3389/fmicb.2018.02000
Kan H, Zhao F, Zhang XX, Ren H, Gao S (2015) Correlations of gut microbial community shift with hepatic damage and growth inhibition of carassius auratus induced by pentachlorophenol exposure. Environ Sci Technol 49:11894–11902. https://doi.org/10.1021/acs.est.5b02990
Kokou F et al (2019) Core gut microbial communities are maintained by beneficial interactions and strain variability in fish. Nature Microbiol 4:2456–2465. https://doi.org/10.1038/s41564-019-0560-0
Kokou F, Sasson G, Nitzan T, Doron-Faigenboim A, Harpaz S, Cnaani A, Mizrahi I (2018) Host genetic selection for cold tolerance shapes microbiome composition and modulates its response to temperature. eLife 7 https://doi.org/10.7554/eLife.36398
Larsen AM, Mohammed HH, Arias CR (2014) Characterization of the gut microbiota of three commercially valuable warmwater fish species. J Appl Microbiol 116:1396–1404. https://doi.org/10.1111/jam.12475
Le D et al (2020) Gut microbiota of migrating wild rabbit fish (Siganus guttatus) larvae have low spatial and temporal variability. Microbial Ecol 79:539–551. https://doi.org/10.1007/s00248-019-01436-1
Li T et al (2017) Bacterial signatures of “red-operculum” disease in the gut of crucian carp (Carassius auratus). Microbial Ecol 74:510–521. https://doi.org/10.1007/s00248-017-0967-1
Li X, Yan Q, Ringø E, Wu X, He Y, Yang D (2016) The influence of weight and gender on intestinal bacterial community of wild largemouth bronze gudgeon (Coreius guichenoti, 1874). BMC Microbiol 16:191. https://doi.org/10.1186/s12866-016-0809-1
Li X, Yu Y, Feng W, Yan Q, Gong Y (2012) Host species as a strong determinant of the intestinal microbiota of fish larvae. J Microbiol (Seoul, Korea) 50:29–37. https://doi.org/10.1007/s12275-012-1340-1
Li X, Zhou L, Yu Y, Ni J, Xu W, Yan Q (2017) Composition of gut microbiota in the Gibel carp (Carassius auratus gibelio) varies with host development. Microbial Ecol 74:239–249. https://doi.org/10.1007/s00248-016-0924-4
Lin SM, Zhou XM, Zhou YL, Kuang WM, Chen YJ, Luo L, Dai FY (2020) Intestinal morphology, immunity and microbiota response to dietary fibers in largemouth bass. Micropterus Salmoide Fish & Shellfish Immunol 103:135–142. https://doi.org/10.1016/j.fsi.2020.04.070
Llewellyn MS et al (2016) The biogeography of the atlantic salmon (Salmo salar) gut microbiome. ISME J 10:1280–1284. https://doi.org/10.1038/ismej.2015.189
Lokesh J, Kiron V, Sipkema D, Fernandes JMO, Moum T (2019) Succession of embryonic and the intestinal bacterial communities of Atlantic salmon (Salmo salar) reveals stage-specific microbial signatures. MicrobiologyOpen 8:e00672. https://doi.org/10.1002/mbo3.672
Mekuchi M, Asakura T, Sakata K, Yamaguchi T, Teruya K, Kikuchi J (2018) Intestinal microbiota composition is altered according to nutritional biorhythms in the leopard coral grouper (Plectropomus leopardus). PloS one 13:e0197256. https://doi.org/10.1371/journal.pone.0197256
Michl SC, Ratten JM, Beyer M, Hasler M, LaRoche J, Schulz C (2017) The malleable gut microbiome of juvenile rainbow trout (Oncorhynchus mykiss): diet-dependent shifts of bacterial community structures. PloS one 12:e0177735. https://doi.org/10.1371/journal.pone.0177735
Mirghaed AT, Yarahmadi P, Hosseinifar SH, Tahmasebi D, Gheisvandi N, Ghaedi A (2018) The effects singular or combined administration of fermentable fiber and probiotic on mucosal immune parameters, digestive enzyme activity, gut microbiota and growth performance of Caspian white fish (Rutilus frisii kutum) fingerlings. Fish & shellfish Immunol 77:194–199. https://doi.org/10.1016/j.fsi.2018.02.007
Navarrete P et al (2012) PCR-TTGE analysis of 16S rRNA from rainbow trout (Oncorhynchus mykiss) gut microbiota reveals host-specific communities of active bacteria. PloS one 7:e31335. https://doi.org/10.1371/journal.pone.0031335
Ni J, Yan Q, Yu Y, Zhang T (2014) Factors influencing the grass carp gut microbiome and its effect on metabolism. FEMS Microbiol Ecol 87:704–714. https://doi.org/10.1111/1574-6941.12256
Parata L, Mazumder D, Sammut J, Egan S (2020) Diet type influences the gut microbiome and nutrient assimilation of genetically improved farmed tilapia (Oreochromis niloticus). PloS one 15:e0237775. https://doi.org/10.1371/journal.pone.0237775
Parshukov AN, Kashinskaya EN, Simonov EP, Hlunov OV, Izvekova GI, Andree KB, Solovyev MM (2019) Variations of the intestinal gut microbiota of farmed rainbow trout, Oncorhynchus mykiss (Walbaum), depending on the infection status of the fish. J App Microbiol 127:379–395. https://doi.org/10.1111/jam.14302
Pérez T, Balcázar JL, Ruiz-Zarzuela I, Halaihel N, Vendrell D, de Blas I, Múzquiz JL (2010) Host-Microbiota Interactions within the Fish Intestinal Ecosystem. Mucosal Immunol 3:355–360. https://doi.org/10.1038/mi.2010.12
Phillips JB, Westerfield M (2014) Zebrafish models in translational research: tipping the scales toward advancements in human health. Dis Model Mech 7:739–743. https://doi.org/10.1242/dmm.015545
Qian H, Zhang M, Liu G, Lu T, Sun L, Pan X (2019) Effects of different concentrations of Microcystis aeruginosa on the intestinal microbiota and immunity of zebrafish (Danio rerio). Chemosphere 214:579–586. https://doi.org/10.1016/j.chemosphere.2018.09.156
Restivo VE, Kidd KA, Surette MG, Servos MR, Wilson JY (2021) Rainbow darter (Etheostoma caeruleum) from a river impacted by municipal wastewater effluents have altered gut content microbiomes. Sci Total Environ 751:141724. https://doi.org/10.1016/j.scitotenv.2020.141724
Robinson CD, Klein HS, Murphy KD, Parthasarathy R, Guillemin K, Bohannan BJM (2018) Experimental bacterial adaptation to the zebrafish gut reveals a primary role for immigration. PLoS Biol 16:e2006893. https://doi.org/10.1371/journal.pbio.2006893
Rolig AS et al (2017) The enteric nervous system promotes intestinal health by constraining microbiota composition. PLoS Biol 15:e2000689. https://doi.org/10.1371/journal.pbio.2000689
Ruiz-Rodríguez M et al (2020) Host species and body site explain the variation in the microbiota associated to wild sympatric Mediterranean teleost fishes. Microb Ecol 80:212–222. https://doi.org/10.1007/s00248-020-01484-y
Schlegel A (2012) Studying non-alcoholic fatty liver disease with zebrafish a confluence of optics, genetics, and physiology. Cell Mol life Sci: CMLS 69:3953–3961. https://doi.org/10.1007/s00018-012-1037-y
Sevellec M, Derome N, Bernatchez L (2018) Holobionts and ecological speciation: the intestinal microbiota of lake whitefish species pairs. Microbiome 6:47. https://doi.org/10.1186/s40168-018-0427-2
Song W et al (2016) The gut microbial community of Antarctic fish detected by 16S rRNA gene sequence analysis. BioMed research international 2016:3241529. https://doi.org/10.1155/2016/3241529
Stagaman K, Burns AR, Guillemin K, Bohannan BJ (2017) The role of adaptive immunity as an ecological filter on the gut microbiota in zebrafish. ISME J 11:1630–1639. https://doi.org/10.1038/ismej.2017.28
Stephens WZ, Burns AR, Stagaman K, Wong S, Rawls JF, Guillemin K, Bohannan BJ (2016) The composition of the zebrafish intestinal microbial community varies across development. ISME J 10:644–654. https://doi.org/10.1038/ismej.2015.140
Sugita H, Kitao S, Narisawa S, Minamishima R, Itoi S (2017) Diversity of culturable bacterial communities in the intestinal tracts of goldfish (Carassius auratus) and their ability to produce N-acyl homoserine lactone. Folia microbiologica 62:263–267. https://doi.org/10.1007/s12223-017-0498-7
Sugita H, Tsunohara M, Ohkoshi T, Deguchi Y (1988) The establishment of an intestinal microflora in developing goldfish (Carassius auratus) of culture ponds. Microbial ecol 15:333–344. https://doi.org/10.1007/bf02012646
Tan HY, Chen SW, Hu SY (2019) Improvements in the growth performance, immunity, disease resistance, and gut microbiota by the probiotic Rummeliibacillus stabekisii in Nile tilapia (Oreochromis niloticus). Fish & shellfish Immunol 92:265–275. https://doi.org/10.1016/j.fsi.2019.06.027
Tran NT, Zhang J, Xiong F, Wang GT, Li WX, Wu SG (2018) Altered gut microbiota associated with intestinal disease in grass carp (Ctenopharyngodon idellus) World. J Microbiol Biotechnol 34:71. https://doi.org/10.1007/s11274-018-2447-2
Trinh LTT, Bakke I, Vadstein O (2017) Correlations of age and growth rate with microbiota composition in Atlantic cod (Gadus morhua). larvae Scientific reports 7:8611. https://doi.org/10.1038/s41598-017-09073-9
Trotter AJ, Parslow AC, Heath JK (2009) Morphologic analysis of the zebrafish digestive system. Methods Mol Biol (Clifton, NJ) 546:289–315. https://doi.org/10.1007/978-1-60327-977-2_18
Wiles TJ et al (2016) Host gut motility promotes competitive exclusion within a model intestinal microbiota. PLoS Biol 14:e1002517. https://doi.org/10.1371/journal.pbio.1002517
Wiles TJ, Schlomann BH, Wall ES, Betancourt R, Parthasarathy R, Guillemin K (2020) Swimming motility of a gut bacterial symbiont promotes resistance to intestinal expulsion and enhances inflammation. PLoS Biol 18:e3000661. https://doi.org/10.1371/journal.pbio.3000661
Xia Y et al (2019) Effects of Lactococcus lactis subsp. lactis JCM5805 on colonization dynamics of gut microbiota and regulation of immunity in early ontogenetic stages of tilapia. Fish & shellfish Immunol 86:53–63. https://doi.org/10.1016/j.fsi.2018.11.022
Xin GY, Li WG, Suman TY, Jia PP, Ma YB, Pei DS (2020) Gut bacteria Vibrio sp. and Aeromonas sp. trigger the expression levels of proinflammatory cytokine: first evidence from the germ-free zebrafish. Fish & shellfish Immunol 106:518–525. https://doi.org/10.1016/j.fsi.2020.08.018
Yang HT, Zou SS, Zhai LJ, Wang Y, Zhang FM, An LG, Yang GW (2017) Pathogen invasion changes the intestinal microbiota composition and induces innate immune responses in the zebrafish intestine. Fish Shellfish Immunol 71:35–42. https://doi.org/10.1016/j.fsi.2017.09.075
Yang S et al (2019) Effects of different diets on the intestinal microbiota and immunity of common carp (Cyprinus carpio). J App Microbiol 127:1327–1338. https://doi.org/10.1111/jam.14405
Yang Y, Tomkovich S, Jobin C (2014) Could a swimming creature inform us on intestinal diseases? Lessons from Zebrafish Inflam Bowel Dis 20:956–966. https://doi.org/10.1097/01.Mib.0000442923.85569.68
Ye L, Mueller O, Bagwell J, Bagnat M, Liddle RA, Rawls JF (2019) High fat diet induces microbiota-dependent silencing of enteroendocrine cells eLife 8 https://doi.org/10.7554/eLife.48479
You C et al (2019) Effects of dietary lipid sources on the intestinal microbiome and health of golden pompano (Trachinotus ovatus). Fish Shellfish Immunol 89:187–197. https://doi.org/10.1016/j.fsi.2019.03.060
Yu J et al (2021) Effects of chronic nitrate exposure on the intestinal morphology, immune status, barrier function, and microbiota of juvenile turbot (Scophthalmus maximus). Ecotoxicol environ safety 207:111287. https://doi.org/10.1016/j.ecoenv.2020.111287
Yuan L, Wang L, Li ZH, Zhang MQ, Shao W, Sheng GP (2019) Antibiotic resistance and microbiota in the gut of Chinese four major freshwater carp from retail markets. Environ Pollution (Barking, Essex: 1987) 255:113327. https://doi.org/10.1016/j.envpol.2019.113327
Zarkasi KZ, Taylor RS, Abell GC, Tamplin ML, Glencross BD, Bowman JP (2016) Atlantic salmon (Salmo salar L.) gastrointestinal microbial community dynamics in relation to digesta properties and diet. Microbial ecol 71:589–603. https://doi.org/10.1007/s00248-015-0728-y
Zhao P, Lu W, Hong Y, Chen J, Dong S, Huang Q (2021) Long-term wet precipitation of PM(2.5) disturbed the gut microbiome and inhibited the growth of marine medaka Oryzias melastigma. Sci Total Environ 755:142512. https://doi.org/10.1016/j.scitotenv.2020.142512
Zhao X, Pack M (2017) Modeling intestinal disorders using zebrafish. Methods Cell Biol 138:241–270. https://doi.org/10.1016/bs.mcb.2016.11.006
Zheng X, Yang R, Hu J, Lin S, Gu Z, Ma Z (2019) The gut microbiota community and antioxidant enzymes activity of barramundi reared at seawater and freshwater. Fish Shellfish Immunol 89:127–131. https://doi.org/10.1016/j.fsi.2019.03.054
Acknowledgements
We sincerely thank Dr. Stanley Li Lin for his careful revision of both language and science. This work was supported by the Natural Science Foundation of China (Grant numbers [81771384]).
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This work was supported by the National Natural Science Foundation of China (Grant numbers [81771384]).
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Yanqin Shen had the idea for the article. Chang Liu and Li-ping Zhao performed the literature search and data analysis. The first draft of the manuscript was written by Chang Liu, and Yanqin Shen critically revised the work. All authors read and approved the final manuscript.
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Liu, C., Zhao, LP. & Shen, YQ. A systematic review of advances in intestinal microflora of fish. Fish Physiol Biochem 47, 2041–2053 (2021). https://doi.org/10.1007/s10695-021-01027-3
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DOI: https://doi.org/10.1007/s10695-021-01027-3