The geometrical origin of the strain-twist coupling in double helices

A simple geometrical explanation for the counterintuitive phenomenon when twist leads to extension in double helices is presented. The coupling between strain and twist is investigated using a tubular description. It is shown that the relation between strain and rotation is universal and depends only on the pitch angle. For pitch angles below 39.4◦ strain leads to further winding, while for larger pitch angles strain leads to unwinding. The zero-twist structure, with a pitch angle of 39.4◦, is at the unique point between winding and unwinding and independent of the mechanical properties of the double helix. The existence of zero-twist structures, i.e. structures that display neither winding, nor unwinding under strain is discussed. Close-packed double helices are shown to extend rather than shorten when twisted. Numerical estimates of this elongation upon winding are given for DNA, chromatin, and RNA.