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GALA quadrupolar wave plots of tilted transmembrane peptides in DLPC bilayers. GWALP23-H12 (red; tilt τ = 23°, rotation ρ = 308°, pH 4) is similar to GWALP23 (black; tilt τ = 21°, rotation ρ = 305°, irrespective of pH), indicating that the buried His12 residue is uncharged. GWALP23-H14 (blue; tilt τ = 27°, rotation ρ = 254°, pH 5.9) is similar to charged GWALP23-R14 (green; tilt τ = 27°, rotation ρ = 260°) (25), suggesting that the His14 residue is charged at pH 5.9.
