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. 2020 Dec 21;8:602994. doi: 10.3389/fcell.2020.602994

Table 3.

Post translational modification of known MARBPs in eukaryotes.

Target MARBP Function References
Phosphorylation
SATB1 PKC Phosphorylates SATB1 leading to increased association with HDAC1 resulting in altered gene expression Ning et al., 2020
Drosophila and mammalian cut homeodomain proteins PKC phosphorylates cut homeodomain repeats 1,2, 3 at Thr415, Thr804, Ser987, respectively, reducing DNA binding ability Han et al., 2013
Murine cut homeodomain proteins CKII phosphorylates cut homeodomain repeats 1, 2, 3 at Ser400, Ser789, Ser972, respectively, decreasing DNA binding ability Coqueret et al., 1996
p110 Cux/CDP Cyclin A/cdk1 phosphorylates p110 Cux/CDP at Ser1237 and Ser1270 in G2 phase of cell cycle resulting in reduced DNA binding activity Coqueret et al., 1998
Lamin B PKCα phosphorylates Lamin B in HL60 cells resulting in proteolysis of Lamin B and subsequent DNA fragmentation Zaremba-Czogalla et al., 2012
GRASS404 sequence of mammalian Lamin C PKC phosphorylates mammalian Lamin C and is important for its nuclear import which then plays a role maintenance of nuclear architecture Eggert et al., 1993; Leukel and Jost, 1995; Shimizu et al., 1998
Chicken Lamin B2 Ser410/Ser411 of Chicken Lamin B2 is phosphorylated by PKC that regulates its import into nucleus that then plays a role in maintenance of nuclear architecture Haas and Jost, 1993
MeCP2 Mutational analysis in Rett's syndrome indicated a role for MeCP2 phosphorylation at Ser80 in modulating its association with chromatin in resting neurons. However, calcium influx causes dephosphorylation causing dissociation. Hennekes et al., 1993
MeCP2 Membrane depolarization caused Ser421 phosphorylation of MeCP2 leading to bdnf promoter activation in neuronal cells by reducing MeCP2 DNA binding activity Tao et al., 2009
AcMFP1 CKII phosphorylates AcMFP1 in a cell cycle dependent manner resulting in reduced nuclear matrix binding Zhou et al., 2006
Chloroplast localized MFP1 in tobacco plants CKII phosphorylates chloroplast MFP1 to decrease association with the chloroplast nucleoid Cohen et al., 2011
HMGA1a Cdc2 phosphorylates Thr52, Thr77, Ser35 of HMGA1a in vitro and in metaphase arrested cells leading to reduced DNA binding ability Nissen et al., 1991; Meier et al., 1996; Samaniego et al., 2006
HMGA1a HIPK2 phosphorylates Thr52, Thr77, Ser35 of HMGA1a leading to reduced DNA binding ability at the human germline ε promoter. Nissen et al., 1991
HMGA1a PKC phosphorylates HMGA1a at Ser43 and Ser63 and attenuates binding with PKC⋎ and neurogranin/RC3 promoter Reeves et al., 1991
HMGA1a Hyperphosphorylation of HMGA1a has been seen in early apoptotic leukemia cells U937, K562, HL60, NB4 leading to displacement from chromatin that helps playing a role in formation of early apoptotic bodies containing condensed chromatin. The ATM kinase pathway plays a role. Zhang and Wang, 2007
HMGA1b ATM kinase phosphorylates at Ser87 within the SQ motif in response to DNA damage and leads to chromosomal reorganization with changes in gene expression. Xiao et al., 2000
Nucleolin Protein Kinase NII phosphorylates nucleolin that leads to increased ribosomal biogenesis in fetal bovine arterial endothelial cells. Diana et al., 2001
SMAR1 SMAR1 is phosphorylated by ATM kinase at Ser370 which helps in recruitment of deacetylated Ku70 at DNA double stranded breaks ensuring timely repair Pentimalli et al., 2008
MUT P53 The N terminus of MUT P53 is phosphorylated by JNK and C-terminus is phosphorylated by PLK2 both leading to enhanced oncogenic activities. Ser392 was phosphorylated in MUT P53 leading to tetramer stabilization and increased DNA binding activity leading to oncogenic properties Ullrich et al., 1993; Minamoto et al., 2001; Matsumoto et al., 2004a
CTCF CTCF is phosphorylated by PLK1 at Ser224 promoting binding at a subset of CTCF binding sites and affects changes in gene expression including upregulation of p21 and p53 during G2/M transition in mouse embryonic stem cell colonies. Matsumoto et al., 2004a
Acetylation
SATB1 Acetylation of SATB1 by PCAF at Lys136 in PDZ domain leads to disruption of interaction with CtBP1 leading to upregulation of target genes. Yap et al., 2004
HMGIY Lys71 is acetylated by PCAF/GCN5 that activates transcription of IFNβ by stabilizing the enhanceosome complex while acetylation at Lys65 by CBP destabilizes this complex leading to downregulation Bode and Dong, 2004
Ku70 An increased Ku70 acetylation in the aggressive form of neuroblastoma N type compared to less aggressive type S reduces the DNA binding ability of Ku70 Luo et al., 2020
MUT P53 Arg273His and Arg248Trp mutants of P53 are hyperacetylated Lys320, Lys373, Lys382 that helps in nuclear accumulation Chaudhary et al., 2014
MUT P53 CBP/P300 mediated acetylation of Lys373 under Id4 overexpression in DU145 cell line promoting DNA binding activity Li et al., 2000
Sumoylation
SATB2 This is sumoylated by PIAS1 at Lys233 and Lys350 downregulating DNA binding activity at immunoglobulin μ gene in pre-B cells. Tang et al., 2008
SAFB1 SAFB1 is SUMO1 labeled by PIAS1 and is important for RNA Pol II recruitment at ribosomal protein and translator genes. Sumoylated SATB1 acts as transcriptional activator. Knowell et al., 2013
SAFB1 Interestingly SAFB1 is modified by SUMO1 and SUMO2/3 at Lys231 and Lys294, respectively. This modification makes it act like a co-repressor activity. Rodriguez et al., 2012
Ku70 Ku70 is Sumoylated at five residues in its C-terminal tail Lys588, Lys591, Lys592, Lys595, and Lys596 thereby promoting DNA binding activity during DNA repair and telomere maintenance Perez et al., 2010
MeCP2 It is sumoylated at Lys223 that is required for transcriptional repression activities. It helps in recruitment of HDAC1/2 complex. In rats abrogating this site leads to poor hippocampal synapse development in rats thus highlighting its importance in central nervous system development. Dobreva et al., 2003
CTCF CTCF is sumoylated by SUMO1, 2, 3 both N-terminal and C-terminal domain by Pc2 E3 ligase. This is important for repression of c-Myc P2 promoter. Liu et al., 2015
CTCF CTCF is desumoylated at Lys74 and 689 by SENP1 under hypoxic conditions in human corneal cells Garee et al., 2011
Methylation
HMGA1a Arg25 of the first AT hook in HMGA1a is found to be methylated in human leukemia cells, rat thyroid tumors cells, human prostate tumor cells during apoptosis. In vivo PRMT1 methylates HMGA1a at this site. Cheng et al., 2014; Hang et al., 2014
HMGB1 In neutrophils HMGB1 is methylated at Lys42 leading to weakening of DNA binding ability causing export to cytoplasm. MacPherson et al., 2009
WTP53 WTP53 is methylated at Lys370 by SMYD2 causes repression of activity while methylation at Lys372 by SET7/9 causes stabilization of chromatin associated SMAR1 Sgarra et al., 2003; Wang et al., 2012
PARylation
PARP1 PARP1 auto-PARylates itself that leads to reduced DNA binding ability Scott et al., 2012
SAFB1 SAFB1 PARylation helps in recruitment of p-⋎-H2AX at damage sites in response to DNA damage responses West and Gozani, 2011
Ku70 Ku70 is PARylated that reduces DNA binding activity and promotes classical NHEJ pathway. D'Amours et al., 1999; Amé et al., 2004; Altmeyer et al., 2013; Morales et al., 2014
DNA-PK catalytic domain PARylation of DNA-PK catalytic subunit increases its kinase activity thereby enhancing its role in double stranded DNA break repair. Mansour et al., 2010; Cheng et al., 2011
HMGI/Y/I-C Incubation of isolated HeLa cell nuclei in calcium containing buffers lead to PARylation of HMGI/Y/I-C proteins leading to probable collapse in nuclear architecture Niedergang et al., 1979; Jones et al., 1989; Boulikas, 1990; Boulikas et al., 1990; Realini and Althaus, 1992; Asher et al., 1995; Belka et al., 1995; Ruscetti et al., 1998; Pleschke et al., 2000; Veuger et al., 2004; Gagne et al., 2008
MeCP2 MeCP2 has lower chromatin binding affinity in PARP-cells therefore substantiating the fact that PARylation of MeCP2 leads to reduced DNA binding affinity Trump and Berezesky, 1995
CTCF PARylated CTCF is enriched at the Cp promoter in type III latency exhibiting EBV immortalized cell line testifying its importance in transcription Miyazaki, 1995
CTCF Studies using human breast 226LDM cells it has been found that several new sites are recognized by CTCF on PARylation Earnshaw, 1995