Purified samples were dispensed on glow-discharged carbon-coated copper grids. Each grid was negatively stained with 2.0% (w/v) uranyl acetate. Micrographs were collected on a direct electron detector (FEI Falcon II) using FEI Tecnai G2 F20 (FEI, Hillsboro, OR, USA) operating at 200 kV. For reference-free 2D class averaging, particle images were boxed out with e2boxer.py in the EMAN2 package (29) and then processed with RELION software (30).

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