Differentiation and dispersal in biofilms

Biofilm formation is now commonly associated with concepts of development, differentiation, and dispersal of microorganisms, and often more broadly with multicellular biological systems. This underlying theme of multicellularity among sessile microorganisms has undoubtedly attracted significant fundamental research interest to the field. This chapter will summarize and discuss aspects of cellular differentiation in biofilms, including microcolony-based dispersal, autolysis of subpopulations of biofilm cells, and the recent finding that nitric oxide—a ubiquitous signal for cellular differentiation—can induce dispersal in Pseudomonas aeruginosa biofilms. Introduction—biofilms as primitive multicellular systems? Recent years have witnessed a dramatic expansion of research into bacterial biofilms, with rapid advances in molecular technologies and microscopy enabling detailed studies of the biofilm mode of life across a range of systems and organisms. This surge of interest is fueled in part by the ubiquitous impact of biofilms, and the need to manipulate, enhance or prevent biofilm formation in diverse environments. In addition, the discovery of cell–cell signaling or quorum sensing systems in bacteria was central in guiding researchers to study bacterial multicellularity and community behavior of bacteria, rather than the previous emphasis on single cell biological processes. Initially, two kinds of multicellular prokaryotic systems were envisaged. One system, as explored for example in differentiation and sporulation in Myxococcus xanthus, involves signaling-mediated development of specialized cells with the ensuing sharing of labor by the different types of cells in the population. The other system refers to the “mob” response displayed by a population of non-differentiating cells, in which the population adopts a new response, such as bioluminescence or virulence factor production, accommodated by the same and simultaneous behavior of all cells. However, since the discovery of a role of quorum sensing in organized surface motility in 1996 (Eberl et al., 1996) and later biofilm development in 1998 (Davies et al., 1998), a third concept has emerged; that of multicellularity and differentiation in biofilms. Biofilms have subsequently been found to display features reminiscent of multicellular systems, including co-ordinated dispersal events, and it now appears that complex development is a hallmark also of the community behavior of classically non-differentiating bacteria (Eberl et al., 1999; Eberl et al., 1996; Klausen et al., 2003; Sauer et al., 2002; Webb et al., 2003b). 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