Floc formation requires cellulose synthase and a specific diguanylate cyclase. (A) Domains of ZM4 proteins with predicted diguanylate cyclase GG(D/E)EF domains (red) or phosphodiesterase EAL domains (yellow). (B) Turbidities of aerobic cultures grown from colonies in a minimal medium (ZYMM) before (filled bars) and after (open bars) cellulase treatment. Shaded circles indicate the same culture pre- and postdispersion. Y, visual observation of flocs; N, no flocs observed. WT, wild-type ZM4; EV, empty vector pSRKT; comp, complementation vector pSRKTc-DGC0919. Error bars, standard deviations. An asterisk indicates a significant difference from the value for the corresponding filled bar (*, P < 0.1) by one-way analysis of variance with Sidak’s multiple-comparison posttest.