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In vitro studies of tissue culture cells with and without surface mucus demonstrated that this increase in phage abundance is mucus dependent and protects the underlying epithelium from bacterial infection. Enrichment of phage in mucus occurs via binding interactions between mucin glycoproteins and Ig-like protein domains exposed on phage capsids. In particular, phage Ig-like domains bind variable glycan residues that coat the mucin glycoprotein component of mucus. Metagenomic analysis found these Ig-like proteins present in the phages sampled from many environments, particularly from locations adjacent to mucosal surfaces. Based on these observations, we present the bacteriophage adherence to mucus model that provides a ubiquitous, but non\u2013host-derived, immunity applicable to mucosal surfaces. The model suggests that metazoan mucosal surfaces and phage coevolve to maintain phage adherence. This benefits the metazoan host by limiting mucosal bacteria, and benefits the phage through more frequent interactions with bacterial hosts. The relationships shown here suggest a symbiotic relationship between phage and metazoan hosts that provides a previously unrecognized antimicrobial defense that actively protects mucosal surfaces.<\/jats:p>","DOI":"10.1073\/pnas.1305923110","type":"journal-article","created":{"date-parts":[[2013,5,21]],"date-time":"2013-05-21T01:03:59Z","timestamp":1369098239000},"page":"10771-10776","update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":924,"title":["Bacteriophage adhering to mucus provide a non\u2013host-derived immunity"],"prefix":"10.1073","volume":"110","author":[{"given":"Jeremy J.","family":"Barr","sequence":"first","affiliation":[{"name":"Department of Biology, San Diego State University, San Diego, CA 92182;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rita","family":"Auro","sequence":"additional","affiliation":[{"name":"Department of Biology, San Diego State University, San Diego, CA 92182;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mike","family":"Furlan","sequence":"additional","affiliation":[{"name":"Department of Biology, San Diego State University, San Diego, CA 92182;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Katrine L.","family":"Whiteson","sequence":"additional","affiliation":[{"name":"Department of Biology, San Diego State University, San Diego, CA 92182;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marcella L.","family":"Erb","sequence":"additional","affiliation":[{"name":"Division of Biological Sciences, University of California, San Diego, CA 92093;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Joe","family":"Pogliano","sequence":"additional","affiliation":[{"name":"Division of Biological Sciences, University of California, San Diego, CA 92093;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Aleksandr","family":"Stotland","sequence":"additional","affiliation":[{"name":"Department of Biology, San Diego State University, San Diego, CA 92182;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Roland","family":"Wolkowicz","sequence":"additional","affiliation":[{"name":"Department of Biology, San Diego State University, San Diego, CA 92182;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andrew S.","family":"Cutting","sequence":"additional","affiliation":[{"name":"Department of Biology, San Diego State University, San Diego, CA 92182;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kelly S.","family":"Doran","sequence":"additional","affiliation":[{"name":"Department of Biology, San Diego State University, San Diego, CA 92182;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Peter","family":"Salamon","sequence":"additional","affiliation":[{"name":"Department of Mathematics and Statistics, San Diego State University, San Diego, CA 92182; and"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Merry","family":"Youle","sequence":"additional","affiliation":[{"name":"Rainbow Rock, Ocean View, HI 96737"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Forest","family":"Rohwer","sequence":"additional","affiliation":[{"name":"Department of Biology, San Diego State University, San Diego, CA 92182;"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"341","published-online":{"date-parts":[[2013,5,20]]},"reference":[{"key":"e_1_3_4_1_2","doi-asserted-by":"crossref","first-page":"1915","DOI":"10.1126\/science.1104816","article-title":"Host-bacterial mutualism in the human intestine","volume":"307","author":"B\u00e4ckhed F","year":"2005","unstructured":"F B\u00e4ckhed, RE Ley, JL Sonnenburg, DA Peterson, JI Gordon, Host-bacterial mutualism in the human intestine. 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