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7, 8, 9 Perturbations of NPCs are known to cause or promote numerous diseases, including viral infections, developmental disorders, neurodegenerative diseases and cancers. The NPC is the largest proteinaceous assembly in vivo, which consists of ~550–1000 molecules of ~30 different nucleoporins (Nups) with the molecular mass of 60–120 MDa from yeast to human, and are the sole gatekeepers controlling the nucleocytoplasmic transport of macromolecules.

4 Since the first glimpse about seventy years ago, 5, 6 NPCs have attracted increasing attentions from researchers. 1, 2, 3 Nuclear pore complexes (NPCs), embedded in NE, are the massive bidirectional transport channels between the nucleus and the cytoplasm.

Macromolecular transport across the two-membraned nuclear envelope (NE) is essential for spatially separated transcription and translation. The presence of nucleus, as the most distinguished hallmark between eukaryotes and prokaryotes, divides a eukaryotic cell into isolated compartments. These structures, together with previously reported structural information of IR, reveal two distinct interaction modes between IR monomers and extensive flexible connections in IR assembly, providing a structural basis for the stability and malleability of IR. In contrast, the intact IR is assembled by loose and instable interactions between IR monomers. In each IR monomer, the Z-shaped Nup188–Nup192 complex in the middle layer is sandwiched by two approximately parallel rhomboidal structures in the inner and outer layers, while Nup188, Nup192 and Nic96 link all subunits to constitute a relatively stable IR monomer. Here we determined the structure of the asymmetric unit of the inner ring (IR monomer) at 3.73 Å resolution by single-particle cryo-electron microscopy, and created an atomic model of the intact IR consisting of 192 molecules of 8 nucleoporins. However, the accurate molecular arrangement of NPCs remains enigmatic owing to their huge size and highly dynamic nature. Nuclear pore complexes (NPCs) mediate bidirectional nucleocytoplasmic transport of substances in eukaryotic cells.

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