  DNA-spools, structures in which DNA is wrapped and helically coiled onto
itself or onto a protein core are ubiquitous in nature. We develop a general
theory describing the non-equilibrium behavior of DNA-spools under linear
tension. Two puzzling and seemingly unrelated recent experimental findings, the
sudden quantized unwrapping of nucleosomes and that of DNA toroidal condensates
under tension are theoretically explained and shown to be of the same origin.
The study provides new insights into nucleosome and chromatin fiber stability
and dynamics.
