[1] These were also rejected due to accumulating experimental evidence from X-ray crystallography, solution NMR, and atomic force microscopy (of both DNA alone, and bound to DNA-binding proteins).
Although localised or transient non-duplex helical structures exist,[2] non-helical models are not currently accepted by the mainstream scientific community.
The most famous of these early models was by Linus Pauling and Robert Corey in 1953 in which they proposed a triple helix with the phosphate backbone on the inside, and the nucleotide bases pointing outwards.
[23] Under torsional stress, a Z-DNA structure can form with opposite twist to B-form DNA, but this is rare within the cellular environment.
[24] The discovery of topoisomerases and gyrases, enzymes that can change the linking number of circular nucleic acids and thus "unwind" and "rewind" the replicating bacterial chromosome, solved the topological objections to the B-form DNA helical structure.
[25] Indeed, in the absence of these topology-altering enzymes, small circular viral and plasmid DNA are inseparable supporting structure whose strands are topologically locked together.