We have characterized and quantified a form of bacterial chemotaxis that manifests only as an emergent property by measuring symmetry breaking in a swarm of Myxococcus xanthus exposed to a two-dimensional nutrient gradient from within an agar substrate. M. xanthus chemotaxis requires cell-cell contact and coordinated motility, as individual motile cells exhibit only nonvectorial movement in the presence of a nutrient gradient. Genes that specifically affect M. xanthus chemotaxis include at least 10 of the 53 that express enhancer binding proteins of the NtrC-like class, an indication that this behavior is controlled through transcription, most likely by a complex signal transduction network.Some of the conditions that define a bacterial swarm-genetically identical cells in close proximity-are theoretically both necessary and sufficient to shift the unit of selection from the individual to the group, i.e., a swarm evolves as a superorganism (5, 34). As a consequence, cellular autonomy diminishes, and each component cell becomes a superposable subunit with a genetic instruction set optimized for the swarm. This instruction set specifies both individual and group responses through interdependent signal transduction networks, which link inputs from neighboring cells and the environment to outputs that control each response (28). The behavior of the swarm evolves to become emergent, manifesting structures, patterns, and properties during the process of self-organization (8). Nature is rife with examples (3, 7).A Myxococcus xanthus swarm is a model organism used to study prokaryotic multicellularity (4, 19). Its best-characterized behavior is the starvation stress response known as development, during which cells break symmetry and move to form dome-shaped aggregates called fruiting bodies (18). Each fruiting body is made up of approximately 1 ϫ 10 5 cells, some of which differentiate into metabolically quiescent and environmentally resistant myxospores (20). From an experimental perspective, a spore-filled fruiting body provides a verifiable endpoint that can be used as a quantifiable metric; a development assay typically measures the number of viable myxospores produced by a swarm over a given period of time following the onset of starvation stress (9). Because these results are quantifiable, they can be used to make definitive statements about relative phenotypic differences among M. xanthus mutant strains, making possible both rank and cluster analysis.Development represents only one response to nutrient limitation, and some researchers have reported a second: M. xanthus can also exhibit a chemotactic response to a gradient of nutrients (22,30). These findings are controversial, however, since other reports claim there is no response (12, 35) (for a summary of the controversy, see Discussion). Unlike development, where sporulation efficiency functions as a quantifiable metric, all previous analyses of M. xanthus swarm chemotaxis have been qualitative in nature. Thus, genetic effects could not be ranked or clustere...