Spontaneous mutation occurs when normal DNA replication takes place after a particularly significant proton has tunnelled. A hydrogen bond joins DNA base pairs. A double well potential along a hydrogen bond separates a potential energy barrier. It is believed that the double well potential is asymmetric, with one well deeper than the other such that the proton normally rests in the deeper well. For a mutation to occur, the proton must have tunnelled into the shallower well. The proton's movement from its regular position is called a tautomeric transition. If DNA replication takes place in this state, the base pairing rule for DNA may be jeopardised, causing a mutation. Per-Olov Lowdin was the first to develop this theory of spontaneous mutation within the double helix. Other instances of quantum tunnelling-induced mutations in biology are believed to be a cause of ageing and cancer.