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The evolutionary emergence of organelles was a defining process in diversifying biochemical reactions within the cell, enabling multi-cellularity. However, compartmentalization also posed a great challenge which is the need to import proteins synthesized in the cytosol into their respective sites of function. While decades of research have set down the fundamental principles of how proteins get from the cytosol into specific organelles, recent studies have brought forward new pathways and additional regulators enabling better definition of the rules governing substrate recognition and targeting. This EMBO Workshop aims for a comprehensive understanding of the molecular mechanism of protein translocation.