Twist-bend instability for toroidal DNA condensates

We propose that semiflexible polymers in poor solvent collapse in two stages. The first stage is the well-known formation of a dense toroidal aggregate. However, if the solvent is sufficiently poor, the condensate will undergo a second structural transition to a twisted entangled state, in which individual filaments lower their bending energy by additionally orbiting around the mean path along which they wind. This topological ripening is consistent with known simulations and experimental results. It connects and rationalizes various experimental observations ranging from strong DNA entanglement in viral capsids to the unusually short pitch of the cholesteric phase of DNA in sperm heads. We propose that topological ripening of DNA toroids could improve the efficiency and stability of gene delivery.