(A) Domain organization of PIKfyve, Fig4, and Vac14. (B) The Vac14 sample used for negative staining EM is pure per SDS-PAGE (line indicates lanes not shown). 2D class averages of Vac14 or MBP-Vac14 show that it pentamerizes. Numbers of particles for each class average are indicated. Maltose binding protein (MBP) fused to the Vac14 N-terminus is at the tip of the Vac14 “leg”. (C) The Fig4/Vac14 sample used for negative staining EM is pure per SDS-PAGE (line indicates lanes not shown). 2D class averages of Fig4/Vac14 show Fig4 between two Vac14 “legs”. (D) Negative stain EM analysis of PIKfyve or MBP+PIKfyve fusions. MBP was inserted into the PIKfyve sequence as indicated by arrows in 1A. The left-most column shows PIKfyve density from the cryo-EM maps (Fig. 2) with docked models of the CCT and kinase modules; a grey ball indicates the location of MBP in the class averages. The other columns show 2D class averages of PIKfyve by itself or with MBP-insertions. (E) The PIKfyve/Fig4/Vac14 sample used for negative staining and cryo-EM is pure per SDS-PAGE. 2D class averages of PIKfyve/Fig4/Vac14 show PIKfyve close to Fig4 at the tip of the Vac14 legs. (F) Schematic overlay of the Vac14 pentamers in Vac14, Fig4/Vac14, and PIKfyve/Fig4/Vac14, showing that the 5-fold symmetry of the Vac14 pentamer is distorted in Fig4/Vac14 and PIKfyve/Fig4/Vac14 complexes.