Early escape mutations confer fitness costs compared to replacement variants. (A) Frequency of variants at amino acid positions 23, 28, and 30 within Gag17-31 in PIC64236. (B) Magnitude of IFN-γ T-cell response by PIC64236 to 15-mers reflecting variation in Gag17-31 at 41 dps (open bars) and 321 dps (filled bars). (C) RK9 epitope (K26/K27/K28 EC50 of 468 ng/ml, black; R28 EC50 of 631 ng/ml, blue; Q26 not recognized, dark gray; Q27 not recognized, light gray) in PIC64236 at 41dps. (D) Highly basic region located in the N-terminal domain of p17. Blue indicates basic residues. The diagram was adapted from reference 53. (E) Fitness costs of observed RK9 epitope variants relative to K28 within an otherwise homologous HIV-1NL4-3 background.