Left–helicase-mediated branch migration (hollow arrow) disassembles a D-loop, followed either by reinvasion or by end annealing to form a NCO. Center, right–CO formation. D-loop branches are captured by ZMM proteins and undergo helicase-mediated branch migration (solid arrows). Branch migration creates a three-arm single-end intermediate (SEI; Hunter and Kleckner, 2001) with a topologically closed bubble (i) or a dHJ (ii). Migration is drawn as unidirectional but may well be reversible. Annealing with the other DSB end and further processing produces a four-armed dHJ that is resolved as a CO. End reinvasion before annealing is also possible but is not drawn here. Center–limited migration can produce a DSBR-like intermediate. Migration after annealing is also possible (Figure S7A), as are different hybrid DNA configurations relative to the CO and DSB (Figure S7B). This model incorporates features of previous models (Allers and Lichten, 2001a; b; Lao et al., 2008).