A hexamer of VCP subunits plus a heterodimer comprising UFDL1 (UFD1) and NPLOC4 bind polyubiquitin that contains lysine-48 linkages (K48polyUb) conjugated to the nascent peptide emerging from the exit tunnel of the 60S ribosomal subunit (inferred from CDC48, the yeast orthologue of VCP, in Brandman et al. 2012, Defenouillère et al. 2013, Verma et al. 2013). In yeast, the NPL4:UFD1 heterodimer (homolog of NPLOC4:UFD1L) acts as an adapter to establish the selectivity of CDC48 binding to K48 polyubiquitin (Tsuchiya et al. 2017, Sato et al. 2019, Williams et al. 2023).
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