Nuclear pore complexes (NPCs) form macromolecular assemblies in the nuclear envelope and mediate bidirectional cargo movement between the nucleus and cytoplasm. Recent evidence suggests that NPCs are not merely transport channels but act as gene regulators. Studies in yeast demonstrate that inducible genes can reposition from the nuclear interior to the nuclear periphery upon activation. At the periphery activated genes engage in physical contacts with nuclear pores. Targeting and tethering of genes to nuclear pores involves multifunctional adaptor complexes, which are thought to couple chromatin modification, transcription and mRNA export. Knowledge of the structure, dynamics and evolution of these adaptor complexes is key to understanding how NPCs control nuclear gene positioning and gene expression. I propose to systematically dissect the architecture and function of NPC-associated adaptor complexes. Our studies will be a unique combination of biochemical and structural approaches in three different model organisms. I anticipate, that this line of research will create a powerful basis to address a number of key questions in the field.
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