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Biofilm cells were resistant to penicillin, but not to rifampin or ciprofloxacin. For some strains, microcolony formation within biofilms was observed. Microcolony formation in strain MC58 depended on a functional copy of the <jats:italic>pilE<\/jats:italic> gene encoding the pilus subunit pilin, and was associated with twitching of cells. Nevertheless, unpiliated <jats:italic>pilE<\/jats:italic> mutants formed biofilms showing that attachment and accumulation of cells did not depend on pilus expression. Mutation and complementation analysis revealed that the type IV pilus\u2010associated protein PilX, which was recently shown to mediate interbacterial aggregation, indirectly supported microcolony formation by contributing to pilus expression. A large number of PilX alleles was identified among genetically diverse meningococcal strains. PilX alleles differed in their propensity to support autoaggregation of cells in suspension, but not in their ability to support microcolony formation within biofilms in the continuous flow system.<\/jats:p>","DOI":"10.1111\/j.1365-2958.2006.05448.x","type":"journal-article","created":{"date-parts":[[2006,10,14]],"date-time":"2006-10-14T20:37:24Z","timestamp":1160858244000},"page":"1292-1309","source":"Crossref","is-referenced-by-count":48,"title":["Meningococcal biofilm formation: structure, development and phenotypes in a standardized continuous flow system"],"prefix":"10.1111","volume":"62","author":[{"given":"Martin","family":"Lappann","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Janus A. 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