1 INTRODUCTION Recent advances in experimental techniques are revealing cellular structure and organization at unprecedented resolution. Cryo-electron tomography (cryo-ET) resolves organelle architectures at nanometer scales (Kühlbrandt, 2022; Maurer et al., 2025; McCafferty et al., 2024; Nogales & Mahamid, 2024), while super-resolution microscopy and spatial proteomics map molecular distributions within cells (Aslam et al., 2017; Guo et al., 2025; Gwosch et al., 2020).