Models were refined against respective cryo-EM maps, which were B factor–sharpened with RELION2’s post-process tool. Automated real-space refinement cycles were alternated with manual model building in Coot. Manual model building was aided by experimental maps filtered at lower resolution and LocScale maps. Geometry statistics were calculated with MolProbity (57). Figures were prepared using PyMOL (58) and Chimera (46).

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