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In Firmicutes, small homodimeric ParA-like (δ 2 ) and ParB-like (ω 2 ) proteins, in concert with cis-acting plasmid-borne  parS  and the host chromosome, secure stable plasmid inheritance in a growing bacterial population. This study shows that (ω:YFP) 2  binding to  parS  facilitates plasmid clustering in the cytosol. (δ:GFP) 2  requires ATP binding but not hydrolysis to localize onto the cell’s nucleoid as a fluorescent cloud. The interaction of (δ:CFP) 2  or δ 2  bound to the nucleoid with (ω:YFP) 2  foci facilitates plasmid capture, from a very broad distribution, towards the nucleoid and plasmid pairing.  parS -bound ω 2  promotes redistribution of (δ:GFP) 2 , leading to the dynamic release of (δ:GFP) 2  from the nucleoid, in a process favored by ATP hydrolysis and protein-protein interaction. (δD60A:GFP) 2 , which binds but cannot hydrolyze ATP, also forms unstable complexes on the nucleoid. In the presence of ω 2 , (δD60A:GFP) 2  accumulates foci or patched structures on the nucleoid. We propose that (δ:GFP) 2  binding to different nucleoid regions and to ω 2 - parS  might generate (δ:GFP) 2  gradients that could direct plasmid movement. The iterative pairing and unpairing cycles may tether plasmids equidistantly on the nucleoid to ensure faithful plasmid segregation by a mechanism compatible with the diffusion-ratchet mechanism as proposed from  in vitro  reconstituted systems.

ParAB Partition Dynamics in Firmicutes: Nucleoid Bound ParA Captures and Tethers ParB-Plasmid Complexes