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Movement of DNA tracking machinery causes topological problems as indicated by the overwinding shown by the circular red arrow. As DNA tracking systems move through the DNA, twists are pushed ahead of replication forks and transcription complexes, resulting in DNA overwinding and the formation of (+)SCs. In the case of replication, precatenanes also form behind the fork; during transcription, (−)SCs form behind the moving DNA tracking machinery. Created with BioRender.com.
