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W. Paerl","year":"1996"},{"issue":"no. 1","key":"ref2","first-page":"34","article-title":"Survival strategies of infectious biofilms","volume":"vol. 13","author":"C. A. Fux","year":"2005"},{"key":"ref3","first-page":"111","article-title":"Mucosal biofilm communities in the human intestinal tract","volume":"vol. 75","author":"S. Macfarlane","year":"2011"},{"issue":"no. 9","key":"ref4","first-page":"453","article-title":"Biofilms as living catalysts in continuous chemical syntheses","volume":"vol. 30","author":"B. Halan","year":"2012"},{"key":"ref5","first-page":"1","article-title":"<italic>Industrial Biofouling<\/italic>: <italic>detection<\/italic>, <italic>prevention and control<\/italic>","author":"J. Jass","year":"2000"},{"issue":"no. 9","key":"ref6","first-page":"623","article-title":"The biofilm matrix","volume":"vol. 8","author":"H. Flemming","year":"2010"},{"issue":"no. 9","key":"ref7","first-page":"563","article-title":"Biofilms: an emergent form of bacterial life","volume":"vol. 14","author":"H.-C. Flemming","year":"2016"},{"issue":"no. 5","key":"ref8","first-page":"222","article-title":"The biofilm matrix\u2014an immobilized but dynamic microbial environment","volume":"vol. 9","author":"I. W. Sutherland","year":"2001"},{"issue":"no. 1","key":"ref9","first-page":"50","article-title":"Solutions to the public goods dilemma in bacterial biofilms","volume":"vol. 24","author":"K. Drescher","year":"2014"},{"issue":"no. 8","key":"ref10","article-title":"A communal bacterial adhesin anchors biofilm and bystander cells to surfaces","volume":"vol. 7","author":"C. Absalon","year":"2011"},{"issue":"no. 4","key":"ref11","first-page":"341","article-title":"Quorum sensing and the social evolution of bacterial virulence","volume":"vol. 19","author":"K. P. Rumbaugh","year":"2009"},{"key":"ref12","first-page":"187","article-title":"Bacterial cellulases and xylanases","volume":"vol. 139","author":"H. J. Gilbert","year":"1993"},{"issue":"no. 4","key":"ref13","first-page":"e0175197","article-title":"Bacterial biofilm under flow : first a physical struggle to stay, then a matter of breathing","volume":"vol. 12","author":"A. Monmeyran","year":"2017"},{"issue":"no. 2","key":"ref14","first-page":"187","article-title":"Mini-review: convection around biofilms","volume":"vol. 28","author":"P. S. Stewart","year":"2012"},{"issue":"no. 4","key":"ref15","first-page":"2175","article-title":"Viscoelasticity of <italic>Staphylococcus aureus<\/italic> biofilms in response to fluid shear allows resistance to detachment and facilitates rolling migration cory","volume":"vol. 71","author":"C. J. Rupp","year":"2005"},{"issue":"no. 9","key":"ref16","first-page":"589","article-title":"Spatial structure, cooperation, and competition in biofilms","volume":"vol. 14","author":"C. D. Nadell","year":"2016"},{"issue":"no. 12","key":"ref17","article-title":"Metabolic and demographic feedbacks shape the emergent spatial structure and function of microbial communities","volume":"vol. 9","author":"S. Estrela","year":"2013"},{"issue":"Suppl, no. Supplement_2","key":"ref18","first-page":"10839","article-title":"Social evolution in multispecies biofilms","volume":"vol. 108","author":"S. Mitri","year":"2011"},{"issue":"no. 2","key":"ref19","first-page":"1","article-title":"Spatial self-organization favors heterotypic cooperation over cheating","volume":"vol. 2013","author":"B. Momeni","year":"2013"},{"issue":"no. 3","key":"ref20","first-page":"876","article-title":"Cooperation and conflict in microbial biofilms","volume":"vol. 104","author":"J. B. 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Kojima","year":"1999"},{"issue":"no. 3","key":"ref42","first-page":"586","article-title":"Steps in the development of a <italic>Vibrio cholerae<\/italic> El Tor biofilm","volume":"vol. 34","author":"P. I. Watnick","year":"1999"},{"issue":"no. 2","key":"ref43","first-page":"400","article-title":"Swimming in circles: motion of bacteria near solid boundaries","volume":"vol. 90","author":"E. Lauga","year":"2006"},{"issue":"no. 3","key":"ref44","first-page":"1","article-title":"Hydrodynamic attraction of swimming microorganisms by surfaces","volume":"vol. 101","author":"A. P. Berke","year":"2008"},{"issue":"no. 6001","key":"ref45","first-page":"197","article-title":"Bacteria use type IV pili to walk upright and detach from surfaces","volume":"vol. 330","author":"M. L. Gibiansky","year":"2010"},{"issue":"no. 6091","key":"ref46","first-page":"236","article-title":"Molecular architecture and assembly principles of <italic>Vibrio cholerae<\/italic> biofilms","volume":"vol. 337","author":"V. 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Steinberg","year":"2002"},{"issue":"no. 110","key":"ref52","first-page":"20150608","article-title":"Competition for space during bacterial colonization of a surface","volume":"vol. 12","author":"D. P. Lloyd","year":"2015"},{"issue":"no. 10","key":"ref53","first-page":"2069","article-title":"Density of founder cells affects spatial pattern formation and cooperation in <italic>Bacillus subtilis<\/italic> biofilms","volume":"vol. 8","author":"J. van Gestel","year":"2014"},{"issue":"no. 1","key":"ref54","first-page":"139","article-title":"Adhesion as a weapon in microbial competition","volume":"vol. 9","author":"J. Schluter","year":"2015"},{"key":"ref55","first-page":"129","article-title":"Optimal foraging theory: the marginal value theorem","volume":"vol. 9","author":"E. L. Charnov","year":"1976"},{"issue":"no. 9","key":"ref56","first-page":"1469","article-title":"Foraging on spatially distributed resources with sub-optimal movement, imperfect information, and travelling costs: departures from the ideal free distribution","volume":"vol. 119","author":"S. Matsumura","year":"2010"},{"issue":"no. 2","key":"ref57","first-page":"383","article-title":"Resource matching with limited knowledge","volume":"vol. 86","author":"E. Ranta","year":"1999"},{"issue":"no. 5","key":"ref58","first-page":"000","article-title":"Perceptual ranges, information gathering, and foraging success in dynamic landscapes","volume":"vol. 189","author":"W. F. Fagan","year":"2017"},{"issue":"no. 3","key":"ref59","first-page":"244","article-title":"Can spatial variation alone lead to selection for dispersal?","volume":"vol. 24","author":"A. Hastings","year":"1983"},{"issue":"no. 2","key":"ref60","first-page":"165","article-title":"Dispersal strategies in patchy environments","volume":"vol. 26","author":"S. A. Levin","year":"1984"},{"issue":"no. 1","key":"ref61","first-page":"17","article-title":"Evolution of dispersal and the ideal free distribution","volume":"vol. 7","author":"R. S. Cantrell","year":"2010"},{"key":"ref62","first-page":"67","article-title":"Microbial bet-hedging: the power of being different","volume":"vol. 25","author":"A. J. Grimbergen","year":"2015"},{"issue":"no. 20","key":"ref63","first-page":"7427","article-title":"Bet-hedging during bacterial diauxic shift","volume":"vol. 111","author":"A. Solopova","year":"2014"},{"issue":"no. 7269","key":"ref64","first-page":"90","article-title":"Experimental evolution of bet hedging","volume":"vol. 462","author":"H. J. E. 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Alexander","year":"1974"},{"issue":"no. 1547","key":"ref70","first-page":"1529","article-title":"Bacteriocins, spite and virulence","volume":"vol. 271","author":"A. Gardner","year":"2004"},{"issue":"no. 6","key":"ref71","first-page":"1195","article-title":"Spite and the scale of competition","volume":"vol. 17","author":"A. Gardner","year":"2004"},{"key":"ref72","first-page":"14371","article-title":"Killing by Type VI secretion drives clonal phase separation and the evolution of cooperation","volume":"vol. 8","author":"L. McNally","year":"2017"},{"issue":"no. 6","key":"ref73","first-page":"E162","article-title":"The Evolution of bacteriocin production in bacterial biofilms","volume":"vol. 178","author":"V. Bucci","year":"2011"},{"issue":"no. 5","key":"ref74","first-page":"1707","article-title":"Demography, altruism, and the benefits of budding","volume":"vol. 19","author":"A. Gardner","year":"2006"},{"issue":"no. 3","key":"ref75","first-page":"318","article-title":"Sex differences in dispersal and the evolution of helping and harming","volume":"vol. 172","author":"R. A. Johnstone","year":"2008"},{"issue":"no. 4","key":"ref76","first-page":"939","article-title":"Limited dispersal, budding dispersal, and cooperation: an experimental study","volume":"vol. 63","author":"R. K\u00fcmmerli","year":"2009"},{"issue":"no. 6","key":"ref77","first-page":"1527","article-title":"The concurrent evolution of cooperation and the population structures that support it","volume":"vol. 65","author":"S. T. Powers","year":"2011"},{"issue":"no. 12","key":"ref78","first-page":"2644","article-title":"The social evolution of dispersal with public goods cooperation","volume":"vol. 26","author":"T. B. 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