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. 2016 May 3;7:73. doi: 10.3389/fgene.2016.00073

Table 1.

CDC48/p97 substrates in the chromatin environment.

Cdc48/p97-dependent process Substrate(s) Cofactor/Accessory factors Experimental system Reference
DNA replication
Replication fork progression, G2/M checkpoint n.d. DVC1 Mammalian cells, patient cells Lessel et al., 2014
Replication licensing CDT-1, CDC-45-GINS UFD-1, NPL-4, UBXN-3/FAF1 C. elegans, X. laevis, mammalian cells Mouysset et al., 2008; Franz et al., 2011, 2016
Replication stress FANCI, FANCD2 DVC1 Mammalian cells Gibbs-Seymour et al., 2015
Replication stress, DNA damage tolerance (DDT) Polη, a.o. DVC1 Mammalian cells, C. elegans Davis et al., 2012; Mosbech et al., 2012
Replication termination MCM7 Dia2 S. cerevisiae, X. laevis Maric et al., 2014; Moreno et al., 2014; Maculins et al., 2015
DNA repair
Diverse genotoxic insults SUMO/Ubiquitin-conjugates Ufd1, Npl4 S. cerevisiae Nie et al., 2012
Diverse genotoxic insults SUMO/Ubiquitin-conjugates Ufd1, Npl4, Rfp1, Pli1 S. pombe Køhler et al., 2013
DNA damage response (DDR) Top1, SUMO-conjugates Wss1, Doa1 S. cerevisiae Balakirev et al., 2015
DNA-double strand break repair L3MBTL1 UFD1, NPL4 Mammalian cells, C. elegans Acs et al., 2011
DNA-double strand break repair SUMO-Rad52 Ufd1 S. cerevisiae, mammalian cells Bergink et al., 2013
DNA-double strand break repair Ubiquitin-(K48)-conjugates UFD1, NPL4 Mammalian cells Meerang et al., 2011
DNA-double strand break repair DNA-PKcs n.d. Mammalian cells Jiang et al., 2013
DNA-protein crosslink (DPC) Top1, a.o. Wss1 S. cerevisiae Stingele et al., 2014
PCNA-dependent response to UV-light n.d. DVC1, mono-ubiquitylated PCNA Mammalian cells Centore et al., 2012
UV-light induced protein turnover CSB UFD1, UBXN7, CUL4 Mammalian cells He et al., 2016
UV-light induced protein turnover CDT1, SET8 UFD1, NPL4, a.o. Mammalian cells, X. leavis Raman et al., 2011
UV-light induced protein turnover DDB2, XPC UFD1, NPL4, UBXN7, CUL4 Mammalian cells Puumalainen et al., 2014
UV-light induced protein turnover Rbp1 Ufd1, Npl4, Ubx4, Ubx5, Cul3 S. cerevisiae Verma et al., 2011
Transcription
Histone ubiquitylation Histone 2B Ubx3 S. cerevisiae, mammalian cells Bonizec et al., 2014
Mating-type switch α2 Ufd1, Npl4, Doa10, a.o. S. cerevisiae Wilcox and Laney, 2009
Transcriptional inactivation HIF1α UBXN7, CUL2, VHL Mammalian cells Alexandru et al., 2008
Transcriptional regulation LexA-VP16, Met4, R-Smads Ufd1, Npl4 S. cerevisiae Ndoja et al., 2014
Heterochromatin decondensation CenH3 Ufd1, Npl4 A. thaliana Merai et al., 2014
Telomere maintenance
Telomerase efficiency Cdc13 Vms1 S. cerevisiae Baek et al., 2012
Telomerase efficiency Est1 Ufd1, Npl4, Ufd4 S. cerevisiae Lin et al., 2015
Sister-chromatid segregation
Anaphase degradation n.d. Ubx4 S. cerevisiae Chien and Chen, 2013
Chromatin decondensation/congression Aurora-B Ufd1, Npl4 X. leavis, mammalian cells Ramadan et al., 2007; Dobrynin et al., 2011
Meiosis AIR-2 n.d. C. elegans Sasagawa et al., 2012
Others
Global analysis SUMO-conjugates Ufd1, STUbL S. pombe Køhler et al., 2015

The table lists the identified chromatin-associated substrates of CDC48/p97, sorted by their functional relevance in indicated cellular processes. In addition, the involvement of cofactors and/or accessory factors is displayed along with the experimental system that was used in the respective publication (a.o. and others, n.d. not determined).