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Negative charge neutralization in the loops and turns reduces hysteresis by 60% at pH 8 relative to WT. (a) Representative titrations illustrating the area in between sigmoidal fits to the data. Hysteresis is reduced further as the number of mutations increase. (b) The area between curves was quantitated from two titrations and standard errors are shown. These data have been normalized by the WT area; numbers are given in Table S3.
