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Small ubiquitin-like modifiers (SUMOs) are a family of small, related proteins (SUMO-1/2/3/4) that can be enzymatically attached to a target protein by a post-translational modification process termed sumoylation, a major regulator of protein function in cellular processes such as nuclear transport, transcriptional regulation, apoptosis and protein stability. This sumoylation is effected by the heterodimeric enzyme SAE1/SAE2 and the SUMO-1-conjugating enzyme Ubch9. The sumoylation pathway mediated by SAE1/SAE2 is distinct from other ubiquitin-like protein (Ubl) pathways.