Fis buffers decrease of negative supercoiling in tyrT and rrnA expression.
2.
Extra helical twists are positive and lead to positive supercoiling, while subtractive twisting causes negative supercoiling.
3.
While topoisomerase IV does relax positive supercoils like DNA gyrase, it does not introduce further negative supercoiling like the latter enzyme.
4.
Negative supercoiling is also thought to favour the transition between B-DNA and Z-DNA, and moderate the interactions of DNA binding proteins involved in gene regulation.