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Based on phase variation frequencies in hpuAB and hmbR , encoding two distinct hemoglobin receptors, Richardson and Stojiljkovic (2001) identified mutL alleles encoding MutL variants with multiple amino acid substitutions in MutL, associated with three distinct switching phenotypes, slow, medium, and fast, together with a mutS allelic variant associated with fast switch and high spontaneous mutation frequency to rifampicin-resistance