Triclosan exposure is associated with rapid restructuring of the microbiome in adult zebrafish

PLoS ONE

Volumn 11, Issue 5, 2016, Pages

  Gaulke, Christopher A ^a   Barton, Carrie L ^a   Proffitt, Sarah ^a   Tanguay, Robert L ^a   Sharpton, Thomas J ^a  

^a OREGON STATE UNIVERSITY   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

RNA 16S; TRICLOSAN; TOPICAL ANTIINFECTIVE AGENT;

ADULT; AEROMONADACEAE; AMPLICON; ARTICLE; COMMUNITY STRUCTURE; CONTROLLED STUDY; ENTEROBACTERIACEAE; ENVIRONMENTAL EXPOSURE; INTESTINE FLORA; MICROBIAL COMMUNITY; MICROBIAL DIVERSITY; MICROBIAL POPULATION DYNAMICS; MICROBIOME RESTRUCTURING; NONHUMAN; ORGANISMAL INTERACTION; POPULATION EXPOSURE; PSEUDOMONAS; RNA SEQUENCE; SEQUENCE ANALYSIS; ZEBRA FISH; ANIMAL; DRUG EFFECTS; DRUG RESISTANCE; GENETICS; METAGENOME; METAGENOMICS; MICROBIOLOGY; MICROFLORA;

ANIMALS; ANTI-INFECTIVE AGENTS, LOCAL; DRUG RESISTANCE; GASTROINTESTINAL MICROBIOME; METAGENOME; METAGENOMICS; MICROBIOTA; RNA, RIBOSOMAL, 16S; TRICLOSAN; ZEBRAFISH;

EID: 84971350797     PISSN: None     EISSN: 19326203     Source Type: Journal    
DOI: 10.1371/journal.pone.0154632     Document Type: Article

References (87)

1. Bohnhoff M, Drake BL, Miller CP. Effect of streptomycin on susceptibility of intestinal tract to experimental Salmonella infection. Exp Biol Med. SAGE Publications; 1954; 86: 132-137.
2. van der Waaij D, Berghuis-de Vries JM, Lekkerkerk Lekkerkerk-v. Colonization resistance of the digestive tract in conventional and antibiotic-treated mice. J Hyg (Lond). 1971; 69: 405-411. doi: 10.1017/S0022172400021653
3. Bouskra D, Brézillon C, Bérard M, Werts C, Varona R, Boneca IG, et al. Lymphoid tissue genesis induced by commensals through NOD1 regulates intestinal homeostasis. Nature. 2008; 456: 507-510. doi: 10.1038/nature07450 PMID: 18987631
4. Mazmanian SK, Round JL, Kasper DL. A microbial symbiosis factor prevents intestinal inflammatory disease. Nature. 2008; 453: 620-625. doi: 10.1038/nature07008 PMID: 18509436
5. Donohoe DR, Wali A, Brylawski BP, Bultman SJ. Microbial Regulation of Glucose Metabolism and Cell-Cycle Progression in Mammalian Colonocytes. PLOS One. 2012; 7. doi: 10.1371/journal.pone.0046589
6. Qin J, Li Y, Cai Z, Li S, Zhu J, Zhang F, et al. A metagenome-wide association study of gut microbiota in type 2 diabetes. Nature. 2012. pp. 55-60. doi: 10.1038/nature11450
7. Koeth RA, Wang Z, Levison BS, Buffa JA, Org E, Sheehy BT, et al. Intestinal microbiota metabolism of L-carnitine, a nutrient in red meat, promotes atherosclerosis. Nat Med. 2013; 19: 576-85. doi: 10.1038/nm.3145 PMID: 23563705
8. Scher JU, Sczesnak A, Longman RS, Segata N, Ubeda C, Bielski C, et al. Expansion of intestinal Prevotella copri correlates with enhanced susceptibility to arthritis. Elife. 2013; 2013. doi: 10.7554/eLife.01202.001
9. Smith MI, Yatsunenko T, Manary MJ, Trehan I, Mkakosya R, Cheng J, et al. Gut microbiomes of Malawian twin pairs discordant for kwashiorkor. Science. 2013; 339: 548-54. doi: 10.1126/science.1229000 PMID: 23363771
10. Dethlefsen L, Relman DA. Incomplete recovery and individualized responses of the human distal gut microbiota to repeated antibiotic perturbation. Proc Natl Acad Sci U S A. 2011; 108 Suppl: 4554-4561. doi: 10.1073/pnas.1000087107
11. Jernberg C, Löfmark S, Edlund C, Jansson JK. Long-term ecological impacts of antibiotic administration on the human intestinal microbiota. ISME J. 2007; 1: 56-66. doi: 10.1038/ismej.2007.3 PMID: 18043614
12. Glavan TW, Gaulke CA, Santos Rocha C, Sankaran-Walters S, Hirao LA, Raffatellu M, et al. Gut immune dysfunction through impaired innate pattern recognition receptor expression and gut microbiota dysbiosis in chronic SIV infection. Mucosal Immunol. Society for Mucosal Immunology; 2015; Available: doi: 10.1038/mi.2015.92
13. Nelson AM, Walk ST, Taube S, Taniuchi M, Houpt ER, Wobus CE, et al. Disruption of the human gut microbiota following Norovirus infection. PLOS One. 2012; 7: e48224. doi: 10.1371/journal.pone.0048224 PMID: 23118957
14. Muegge BD, Kuczynski J, Knights D, Clemente JC, González A, Fontana L, et al. Diet drives convergence in gut microbiome functions across mammalian phylogeny and within humans. Science. 2011; 332: 970-974. doi: 10.1126/science.1198719 PMID: 21596990
15. David LA, Maurice CF, Carmody RN, Gootenberg DB, Button JE, Wolfe BE, et al. Diet rapidly and reproducibly alters the human gut microbiome. Nature. 2014; 505: 559-63. doi: 10.1038/nature12820 PMID: 24336217
16. Rappaport SM, Barupal DK, Wishart D, Vineis P, Scalbert A. The blood exposome and its role in discovering causes of disease. Environ Health Perspect. 2014; 122: 769-774. doi: 10.1289/ehp.1308015 PMID: 24659601
17. Wild CP, Scalbert A, Herceg Z. Measuring the exposome: a powerful basis for evaluating environmental exposures and cancer risk. Environ Mol Mutagen. 2013; 54: 480-99. doi: 10.1002/em.21777 PMID: 23681765
18. Nicholson JK, Holmes E, Kinross J, Burcelin R, Gibson G, Jia W, et al. Host-gut microbiota metabolic interactions. Science (80-). 2012; 336: 1262-7. doi: 10.1126/science.1223813
19. Lu K, Abo RP, Schlieper KA, Graffam ME, Levine S, Wishnok JS, et al. Arsenic exposure perturbs the gut microbiome and its metabolic profile in mice: An integrated metagenomics and metabolomics analysis. Environ Health Perspect. 2014; 122: 284-291. doi: 10.1289/ehp.1307429 PMID: 24413286
20. Breton J, Massart S, Vandamme P, De Brandt E, Pot B, Foligné B. Ecotoxicology inside the gut: impact of heavy metals on the mouse microbiome. BMC Pharmacol Toxicol. 2013; 14: 62. doi: 10.1186/2050-6511-14-62 PMID: 24325943
21. Liu Y, Li Y, Liu K, Shen J. Exposing to cadmium stress cause profound toxic effect on microbiota of the mice intestinal tract. PLOS One. 2014; 9. doi: 10.1371/journal.pone.0085323
22. Pinto PIS, Guerreiro EM, Power DM. Triclosan interferes with the thyroid axis in the zebrafish (Danio rerio). Toxicology Research. 2013. doi: 10.1039/c2tx20005h
23. Darbre PD, Harvey PW. Paraben esters: Review of recent studies of endocrine toxicity, absorption, esterase and human exposure, and discussion of potential human health risks. Journal of Applied Toxicology. 2008. pp. 561-578. doi: 10.1002/jat.1358 PMID: 18484575
24. Magee EA, Edmond LM, Tasker SM, Kong SC, Curno R, Cummings JH. Associations between diet and disease activity in ulcerative colitis patients using a novel method of data analysis. Nutr J. 2005; 4: 7. doi: 10.1186/1475-2891-4-7 PMID: 15705205
25. Spitsbergen J, Kent M. The State of the Art of the Zebrafish Model for Toxicology and Toxicologic Pathology Research - Advantages and Current Limitations. Toxicol Pathol. 2003; 31: 62-87. doi: 10.1080/01926230390174959 PMID: 12597434
26. Kent ML, Buchner C, Barton C, Tanguay RL. Toxicity of chlorine to zebrafish embryos. Dis Aquat Organ. 2014; 107: 235-40. doi: 10.3354/dao02683 PMID: 24429474
27. Truong L, Denardo MA, Kundu S, Collins TJ, Tanguay RL. Zebrafish Assays as Developmental Toxicity Indicators in The Design of TAML Oxidation Catalysts. Green Chem. 2013; 15: 2339-2343. doi: 10.1039/C3GC40376A PMID: 24748850
28. Wienholds E, van Eeden F, Kosters M, Mudde J, Plasterk RHA, Cuppen E. Efficient target-selected mutagenesis in zebrafish. Genome Res. 2003; 13: 2700-2707. doi: 10.1101/gr.1725103 PMID: 14613981
29. Grunwald DJ, Eisen JS. Headwaters of the zebrafish - emergence of a new model vertebrate. Nat Rev Genet. 2002; 3: 717-724. PMID: 12209146
30. Burns CG, Milan DJ, Grande EJ, Rottbauer W, MacRae CA, Fishman MC. High-throughput assay for small molecules that modulate zebrafish embryonic heart rate. Nat Chem Biol. 2005; 1: 263-264. doi: 10.1038/nchembio732 PMID: 16408054
31. Truong L, Mandrell D, Mandrell R, Simonich M, Tanguay RL. A rapid throughput approach identifies cognitive deficits in adult zebrafish from developmental exposure to polybrominated flame retardants. Neurotoxicology. 2014; 43: 134-42. doi: 10.1016/j.neuro.2014.03.005 PMID: 24674958
32. Stephens WZ, Wiles TJ, Martinez ES, Jemielita M, Burns AR, Parthasarathy R, et al. Identification of Population Bottlenecks and Colonization Factors during Assembly of Bacterial Communities within the Zebrafish Intestine. MBio. 2015; 6: e01163-15. doi: 10.1128/mBio.01163-15 PMID: 26507229
33. Jemielita M, Taormina MJ, Burns AR, Hampton JS, Rolig AS, Guillemin K, et al. Spatial and Temporal Features of the Growth of a Bacterial Species Colonizing the Zebrafish Gut. MBio. 2014; 5: 1-8. doi: 10.1128/mBio.01751-14
34. Zac Stephens W, Burns AR, Stagaman K, Wong S, Rawls JF, Guillemin K, et al. The composition of the zebrafish intestinal microbial community varies across development. ISME J. International Society for Microbial Ecology; 2016. pp. 644-654. Available: doi: 10.1038/ismej.2015.140
35. Rolig AS, Parthasarathy R, Burns AR, Bohannan BJM, Guillemin K. Individual Members of the Microbiota Disproportionately Modulate Host Innate Immune Responses. Cell Host Microbe. Elsevier; 2015; 18: 613-620.
36. Kanther M, Tomkovich S, Xiaolun S, Grosser MR, Koo J, Flynn EJ, et al. Commensal microbiota stimulate systemic neutrophil migration through induction of Serum amyloid A. Cell Microbiol. 2014; 16: 1053-1067. doi: 10.1111/cmi.12257 PMID: 24373309
37. Semova I, Carten JD, Stombaugh J, Mackey LC, Knight R, Farber SA, et al. Microbiota regulate intestinal absorption and metabolism of fatty acids in the zebrafish. Cell Host Microbe. 2012; 12: 277-88. doi: 10.1016/j.chom.2012.08.003 PMID: 22980325
38. Schweizer HP. Triclosan: A widely used biocide and its link to antibiotics. FEMS Microbiology Letters. 2001. pp. 1-7. doi: 10.1016/S0378-1097(01)00273-7
39. Sandborgh-Englund G, Adolfsson-Erici M, Odham G, Ekstrand J. Pharmacokinetics of triclosan following oral ingestion in humans. Journal of toxicology and environmental health. Part A. 2006. doi: 10.1080/15287390600631706
40. Moss T, Howes D, Williams FM. Percutaneous penetration and dermal metabolism of triclosan (2,4, 4′-trichloro-2′-hydroxydiphenyl ether). Food Chem Toxicol. 2000; 38: 361-370. PMID: 10722890
41. Allmyr M, Adolfsson-Erici M, McLachlan MS, Sandborgh-Englund G. Triclosan in plasma and milk from Swedish nursing mothers and their exposure via personal care products. Sci Total Environ. 2006; 372: 87-93. doi: 10.1016/j.scitotenv.2006.08.007 PMID: 17007908
42. Stoker TE, Gibson EK, Zorrilla LM. Triclosan exposure modulates estrogen-dependent responses in the female wistar rat. Toxicol Sci. 2010; 117: 45-53. doi: 10.1093/toxsci/kfq180 PMID: 20562219
43. Yueh M-F, Taniguchi K, Chen S, Evans RM, Hammock BD, Karin M, et al. The commonly used antimicrobial additive triclosan is a liver tumor promoter. Proc Natl Acad Sci U S A. 2014; doi: 10.1073/pnas.1419119111
44. Wallet MA, Calderon NL, Alonso TR, Choe CS, Catalfamo DL, Lalane CJ, et al. Triclosan alters antimicrobial and inflammatory responses of epithelial cells. Oral Dis. 2013; 19: 296-302. doi: 10.1111/odi.12001 PMID: 24079913
45. Harrow DI, Felker JM, Baker KH. Impacts of triclosan in greywater on soil microorganisms. Appl Environ Soil Sci. Hindawi Publishing Corporation; 2011; 2011.
46. Nietch CT, Quinlan EL, Lazorchak JM, Impellitteri CA, Raikow D, Walters D. Effects of a chronic lower range of triclosan exposure on a stream mesocosm community. Environ Toxicol Chem. 2013; 32: 2874-2887. doi: 10.1002/etc.2385 PMID: 24038532
47. Narrowe AB, Albuthi-Lantz M, Smith EP, Bower KJ, Roane TM, Vajda AM, et al. Perturbation and restoration of the fathead minnow gut microbiome after low-level triclosan exposure. Microbiome. BioMed Central Ltd; 2015; 3: 6.
48. Caporaso JG, Lauber CL, Walters WA, Berg-Lyons D, Huntley J, Fierer N, et al. Ultra-high-throughput microbial community analysis on the Illumina HiSeq and MiSeq platforms. The ISME Journal. 2012. pp. 1621-1624. doi: 10.1038/ismej.2012.8 PMID: 22402401
49. Caporaso JG, Lauber CL, Walters WA, Berg-Lyons D, Lozupone CA, Turnbaugh PJ, et al. Global patterns of 16S rRNA diversity at a depth of millions of sequences per sample. Proc Natl Acad Sci U S A. 2011; 108 Suppl: 4516-4522. doi: 10.1073/pnas.1000080107
50. Caporaso JG, Kuczynski J, Stombaugh J, Bittinger K, Bushman FD, Costello EK, et al. QIIME allows analysis of high-throughput community sequencing data. Nature methods. 2010. pp. 335-336. doi: 10.1038/nmeth.f.303 PMID: 20383131
51. Storey JD. A direct approach to false discovery rates. J R Stat Soc Ser B (Statistical Methodol. Wiley Online Library; 2002; 64: 479-498.
52. Roeselers G, Mittge EK, Stephens WZ, Parichy DM, Cavanaugh CM, Guillemin K, et al. Evidence for a core gut microbiota in the zebrafish. The ISME Journal. 2011. pp. 1595-1608. doi: 10.1038/ismej.2011.38 PMID: 21472014
53. Llewellyn MS, Boutin S, Hoseinifar SH, Derome N. Teleost microbiomes: the state of the art in their characterization, manipulation and importance in aquaculture and fisheries. Front Microbiol. 2014; 5: 207. doi: 10.3389/fmicb.2014.00207 PMID: 24917852
54. Heath RJ, Rubin JR, Holland DR, Zhang E, Snow ME, Rock CO. Mechanism of triclosan inhibition of bacterial fatty acid synthesis. J Biol Chem. 1999; 274: 11110-11114. doi: 10.1074/jbc.274.16.11110 PMID: 10196195
55. Kim YM, Murugesan K, Schmidt S, Bokare V, Jeon JR, Kim EJ, et al. Triclosan susceptibility and cometabolism - A comparison for three aerobic pollutant-degrading bacteria. Bioresour Technol. 2011; 102: 2206-2212. doi: 10.1016/j.biortech.2010.10.009 PMID: 21041079
56. Chuanchuen R, Beinlich K, Hoang TT, Becher A, Karkhoff-Schweizer RR, Schweizer HP. Cross-resistance between triclosan and antibiotics in Pseudomonas aeruginosa is mediated by multidrug efflux pumps: Exposure of a susceptible mutant strain to triclosan selects nfxB mutants overexpressing MexCD-OprJ. Antimicrob Agents Chemother. 2001; 45: 428-432. doi: 10.1128/AAC.45.2.428-432.2001 PMID: 11158736
57. McMurry LM, Oethinger M, Levy SB. Triclosan targets lipid synthesis. Nature. 1998. pp. 531-532. doi: 10.1038/28970
58. Zhu L, Lin J, Ma J, Cronan JE, Wang H. Triclosan resistance of Pseudomonas aeruginosa PAO1 is due to FabV, a triclosan-resistant enoyl-acyl carrier protein reductase. Antimicrob Agents Chemother. 2010; 54: 689-698. doi: 10.1128/AAC.01152-09 PMID: 19933806
59. Lee IH, Kim EJ, Cho YH, Lee JK. Characterization of a novel enoyl-acyl carrier protein reductase of diazaborine-resistant Rhodobacter sphaeroides mutant. Biochem Biophys Res Commun. 2002; 299: 621-627. doi: 10.1016/S0006-291X(02)02702-X PMID: 12459184
60. Khan R, Lee MH, Joo H, Jung Y-H, Ahmad S, Choi J, et al. Triclosan Resistance in a Bacterial Fish Pathogen Aeromonas salmonicida subsp. salmonicida is Mediated by an Enoyl Reductase FabV. J Microbiol Biotechnol. 2015; 25: 511-520. doi: 10.4014/jmb.1407.07021 PMID: 25370725
61. Dufrêne M, Legendre P. Species assemblages and indicator species: the need for a flexible asymmetrical approach. Ecol Monogr. 1997; 67: 345-366. doi: 10.1890/0012-9615(1997)067[0345:SAAIST]2.0.CO;2
62. Arumugam M, Raes J, Pelletier E, Le Paslier D, Yamada T, Mende DR, et al. Enterotypes of the human gut microbiome. Nature. 2011; 473: 174-180. doi: 10.1038/nature10187 PMID: 21508958
63. Endesfelder D, Castell WZ, Ardissone A, Davis-Richardson AG, Achenbach P, Hagen M, et al. Compromised gut microbiota networks in children with anti-islet cell autoimmunity. Diabetes. 2014; 63: 2006-2014. doi: 10.2337/db13-1676 PMID: 24608442
64. Coyte KZ, Schluter J, Foster KR. The ecology of the microbiome: Networks, competition, and stability. Science. 2015; 350: 663-666. doi: 10.1126/science.aad2602 PMID: 26542567
65. Foster KR, Bell T. Competition, not cooperation, dominates interactions among culturable microbial species. Curr Biol. 2012; 22: 1845-1850. doi: 10.1016/j.cub.2012.08.005 PMID: 22959348
66. Newman M. Networks an Introduction [Internet]. Book. Oxford: Oxford University Press; 2010. doi: 10.1093/acprof:oso/9780199206650.001.0001
67. Wilkinson DM, Huberman BA. A method for finding communities of related genes. Proc Natl Acad Sci U S A. 2004; 101 Suppl: 5241-5248. doi: 10.1073/pnas.0307740100
68. Rives AW, Galitski T. Modular organization of cellular networks. Proc Natl Acad Sci USA. 2003; 100: 1128-1133. doi: 10.1073/pnas.0237338100 PMID: 12538875
69. Lewis ACF, Jones NS, Porter MA, Deane CM. The function of communities in protein interaction networks at multiple scales. BMC Syst Biol. 2010; 4: 100. doi: 10.1186/1752-0509-4-100 PMID: 20649971
70. Faust K, Raes J. Microbial interactions: from networks to models. Nat Rev Microbiol. 2012; 10: 538-550. doi: 10.1038/nrmicro2832 PMID: 22796884
71. Yazdankhah SP, Scheie AA, Høiby EA, Lunestad B-T, Heir E, Fotland TØ, et al. Triclosan and antimicrobial resistance in bacteria: an overview. Microb Drug Resist. 2006; 12: 83-90. doi: 10.1089/mdr.2006.12.83 PMID: 16922622
72. Cho I, Yamanishi S, Cox L, Methé BA, Zavadil J, Li K, et al. Antibiotics in early life alter the murine colonic microbiome and adiposity. Nature. 2012; 488: 621-626. doi: 10.1038/nature11400 PMID: 22914093
73. Loo VG, Bourgault A-M, Poirier L, Lamothe F, Michaud S, Turgeon N, et al. Host and pathogen factors for Clostridium difficile infection and colonization. N Engl J Med. 2011; 365: 1693-703. doi: 10.1056/NEJMoa1012413 PMID: 22047560
74. Thomas C, Stevenson M, Riley T V. Antibiotics and hospital-acquired Clostridium difficile-associated diarrhoea: A systematic review. J Antimicrob Chemother. 2003; 51: 1339-1350. doi: 10.1093/jac/dkg254 PMID: 12746372
75. Drury B, Scott J, Rosi-Marshall EJ, Kelly JJ. Triclosan exposure increases triclosan resistance and influences taxonomic composition of benthic bacterial communities. Environ Sci Technol. 2013; 47: 8923-8930. doi: 10.1021/es401919k PMID: 23865377
76. Meade MJ, Waddell RL, Callahan TM. Soil bacteria Pseudomonas putida and Alcaligenes xylosoxidans subsp. denitrificans inactivate triclosan in liquid and solid substrates. FEMS Microbiol Lett. 2001; 204: 45-48. doi: 10.1016/S0378-1097(01)00377-9 PMID: 11682176
77. Jones GL, Russell a. D, Caliskan Z, Stickler DJ. A strategy for the control of catheter blockage by crystalline Proteus mirabilis biofilm using the antibacterial agent triclosan. Eur Urol. 2005; 48: 838-845. doi: 10.1016/j.eururo.2005.07.004 PMID: 16126323
78. Schaffner DW, Bowman JP, English DJ, Fischler GE, Fuls JL, Krowka JF, et al. Quantitative microbial risk assessment of antibacterial hand hygiene products on risk of shigellosis. J Food Prot. 2014; 77: 574-82. doi: 10.4315/0362-028X.JFP-13-366 PMID: 24680068
79. Birosová L, Mikulásová M. Development of triclosan and antibiotic resistance in Salmonella enterica serovar Typhimurium. J Med Microbiol. 2009; 58: 436-441. doi: 10.1099/jmm.0.003657-0 PMID: 19273638
80. Ciusa ML, Furi L, Knight D, Decorosi F, Fondi M, Raggi C, et al. A novel resistance mechanism to triclosan that suggests horizontal gene transfer and demonstrates a potential selective pressure for reduced biocide susceptibility in clinical strains of Staphylococcus aureus. Int J Antimicrob Agents. 2012; 40: 210-220. doi: 10.1016/j.ijantimicag.2012.04.021 PMID: 22789727
81. Mutlu EA, Keshavarzian A, Losurdo J, Swanson G, Siewe B, Forsyth C, et al. A compositional look at the human gastrointestinal microbiome and immune activation parameters in HIV infected subjects. PLOS Pathog. 2014; 10: e1003829. doi: 10.1371/journal.ppat.1003829 PMID: 24586144
82. Galimanas V, Hall MW, Singh N, Lynch MDJ, Goldberg M, Tenenbaum H, et al. Bacterial community composition of chronic periodontitis and novel oral sampling sites for detecting disease indicators. Microbiome. 2014; 2: 32. doi: 10.1186/2049-2618-2-32 PMID: 25225610
83. Yildirim S, Yeoman CJ, Janga SC, Thomas SM, Ho M, Leigh SR, et al. Primate vaginal microbiomes exhibit species specificity without universal Lactobacillus dominance. ISME J. 2014; 8: 2431-44. doi: 10.1038/ismej.2014.90 PMID: 25036926
84. Kembel SW, Jones E, Kline J, Northcutt D, Stenson J, Womack AM, et al. Architectural design influences the diversity and structure of the built environment microbiome. ISME J. 2012; 6: 1469-79. doi: 10.1038/ismej.2011.211 PMID: 22278670
85. The Human Microbiome Project Consortium. Structure, function and diversity of the healthy human microbiome. Nature. 2012; 486: 207-14. doi: 10.1038/nature11234 PMID: 22699609
86. Rawls JF, Mahowald MA, Ley RE, Gordon JI. Reciprocal Gut Microbiota Transplants from Zebrafish and Mice to Germ-free Recipients Reveal Host Habitat Selection. Cell. 2006; 127: 423-433. doi: 10.1016/j.cell.2006.08.043 PMID: 17055441
87. Fortunato S. Community detection in graphs. Physics Reports. 2010. pp. 75-174. doi: 10.1016/j.physrep.2009.11.002