Successful segregation of newly replicated DNA requires that all interlinks between the two strands of duplex DNA be removed, as was pointed out when the structure of DNA was determined In the Escherichia coli chromosome, ∼450,000 links need to be removed each generation, with the two type II topoisomerases, DNA gyrase and topoisomerase IV (TopoIV) being responsible for most of this unlinking.
DNA gyrase acts preferentially to remove positive supercoiling that accumulates ahead of the replication fork, while TopoIV acts potentially both in front of and behind replication forks as they proceed. Inactivation of TopoIV has little or no effect on growth or DNA replication rate, but prevents segregation of newly replicated sister loci and thus cell division, thereby demonstrating its essential role in decatenation.