Knob densities on IEs were measured by AFM essentially as described by Quadt et al. (7 (link)). AFM images were captured either with an MFP-3D microscope (Asylum Research, Santa Barbara, CA, USA) or a Multimode8 AFM (Bruker Nano Inc., Santa Barbara, CA, USA). We used copper grids to locate the IEs via an integrated optical microscope. Images were captured in air under ambient conditions with tapping mode and using a silicon microcantilever (OMCL-AC160TS-W2; Olympus) with a spring constant of 42 N/m and a resonant frequency of ~300 kHz. The images were 512 by 512 pixels and captured at scan speeds of 0.5 to 2.0 Hz, depending on the scan size (0.25 to 15 µm). Scan speeds were optimized individually to minimize noise and integral and proportional gains.