Similar “above-the-bridge-helix” translocation intermediates captured in pausing/arrested Pol II ECs and a common 5caC-recognition mode shared by a variety of 5caC-recognition proteins. a–c. Superimposition of 5caC-paused Pol II EC with CPD-lesion-arrested EC (PDB: 4A93) (a), pyriplatin-lesion-arrested EC (PDB: 3M4O) (b), and α-amanitin-arrested EC (PDB: 2VUM) (c), respectively. The similar “above-the-bridge-helix” translocation intermediates region for accommodation of i+1 5caC (yellow) and DNA lesion (or translocation intermediate captured by α-amanitin) (blutewhite) is highlighted by a red-dotted oval. The damage-arrested or α-amanitin-arrested Pol II ECs are shown in gray. d–f, The conserved interactions and residue involved 5caC recognition by Pol II (Rpb 2-Q531) (d), Wilms tumor protein 1 (Q369, PDB: 4R2R) (e), and human thymine DNA glycosylase (N157, PDB: 3UO7) (f).