Chargaff (1950) discovered the equivalence rule, which suggested that despite wide compositional variations exhibited by different types of DNA, the total amount of purines equals to the amount of pyrimidine (A+G=T+C). The amount of adenine equals to the amount of thymine (A=T) and the amount of guanine equals to the amount of cytosine (G=C). In contrast, the (T+A/G+ C) ratio varied widely in DNAs of different species. In case of double-stranded DNA (A+G)/(C+T)C/G and (G+T)/(A+C) ratios are always or almost equal to one.