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== In the environment == {{Main|Marine bacteriophage}} [[Metagenomics]] has allowed the in-water detection of bacteriophages that was not possible previously.<ref name="pmid12384570">{{cite journal | vauthors = Breitbart M, Salamon P, Andresen B, Mahaffy JM, Segall AM, Mead D, Azam F, Rohwer F | title = Genomic analysis of uncultured marine viral communities | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 99 | issue = 22 | pages = 14250–14255 | date = October 2002 | pmid = 12384570 | pmc = 137870 | doi = 10.1073/pnas.202488399 | doi-access = free | bibcode = 2002PNAS...9914250B | author-link1 = Mya Breitbart }}</ref> Also, bacteriophages have been used in [[hydrology|hydrological]] tracing and modelling in [[river]] systems, especially where surface water and [[groundwater]] interactions occur. The use of phages is preferred to the more conventional [[dye]] marker because they are significantly less absorbed when passing through ground waters and they are readily detected at very low concentrations.<ref>{{cite journal | vauthors = Martin C | doi = 10.1111/j.1747-6593.1988.tb01352.x | title = The Application of Bacteriophage Tracer Techniques in South West Water | journal = Water and Environment Journal | volume = 2 | issue = 6 | pages = 638–642 | year = 1988 | bibcode = 1988WaEnJ...2..638M }}</ref> Non-polluted water may contain approximately 2×10<sup>8</sup> bacteriophages per ml.<ref>{{cite journal | vauthors = Bergh O, Børsheim KY, Bratbak G, Heldal M | title = High abundance of viruses found in aquatic environments | journal = Nature | volume = 340 | issue = 6233 | pages = 467–468 | date = August 1989 | pmid = 2755508 | doi = 10.1038/340467a0 | s2cid = 4271861 | bibcode = 1989Natur.340..467B }}</ref> Bacteriophages are thought to contribute extensively to [[horizontal gene transfer]] in natural environments, principally via [[transduction (genetics)|transduction]], but also via [[transformation (genetics)|transformation]].<ref name="KeenBliskovsky2017">{{cite journal | vauthors = Keen EC, Bliskovsky VV, Malagon F, Baker JD, Prince JS, Klaus JS, Adhya SL | title = Novel "Superspreader" Bacteriophages Promote Horizontal Gene Transfer by Transformation | journal = mBio | volume = 8 | issue = 1 | pages = e02115–16 | date = January 2017 | pmid = 28096488 | pmc = 5241400 | doi = 10.1128/mBio.02115-16 }}</ref> Metagenomics-based studies also have revealed that [[virome]]s from a variety of environments harbor antibiotic-resistance genes, including those that could confer [[multidrug resistance]].<ref name="Lekunberri2017">{{cite journal | vauthors = Lekunberri I, Subirats J, Borrego CM, Balcázar JL | title = Exploring the contribution of bacteriophages to antibiotic resistance | journal = Environmental Pollution | volume = 220 | issue = Pt B | pages = 981–984 | date = January 2017 | pmid = 27890586 | doi = 10.1016/j.envpol.2016.11.059 | bibcode = 2017EPoll.220..981L | hdl = 10256/14115 }}</ref> Recent findings have mapped the complex and intertwined arsenal of anti-phage defense tools in environmental bacteria.<ref>{{cite journal | vauthors = Beavogui A, Lacroix A, Wiart N, Poulain J, Delmont TO, Paoli L, Wincker P, Oliveira PH | title = The defensome of complex bacterial communities | journal = Nature Communications | volume = 15 | issue = 1 | page = 2146 | date = March 2024 | pmid = 38459056 | pmc = 10924106 | doi = 10.1038/s41467-024-46489-0 | bibcode = 2024NatCo..15.2146B }}</ref>
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