An observed transition-transversion ratio compares the number of nucleotide transitions with the number of nucleotide transversions in a specified set of sequence alignment differences:Six transition differences and two transversion differences give . If no transversion is observed, this ordinary finite ratio is undefined; the raw counts should remain explicit.
The instantaneous rate parameter can use a different convention. In the Kimura two-parameter model, let be the rate to the one transition target and the rate to each of the two transversion targets from a base. ThenThus corresponds to twice as many instantaneous transition events as total transversion events, not four times as many.[ref]
A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences by Motoo Kimura. Two-rate substitution model; per-target transition and transversion definitions
Observed endpoint differences can hide repeated nucleotide substitutions. Under independent uniform endpoint bases, the four directed transition mismatches and eight transversion mismatches give a limiting ratio , even when short-time event rates differ. Always state whether the ratio describes scored differences, total events or rates per possible target.
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