Table 1.

Modifications of YB proteins and probable effects on their functions. The complete list of detected modifications can be found at www.phosphosite.org. The names of respective YB proteins except YB-1 (default) are shown in bold in parentheses. Notation: ↑—up-regulation; ↓—down-regulation; n/e—no effect; ?—indirect evidence; ??—unknown function.

Modification	Location	Enzyme	Probable Effect	Reference
Acetylation
Lys81	CSD	HDAC1 (?) HDAC3 (?)	YB-1 binding and translational activation of G3BP1, NFE2L2, HIF-1α mRNAs (↓)	[125]
Lys 303, Lys304	CTD	??	YB-1 secretion (↑)	[153,154]
O-GlcNAcylation
Ser32	A/P	OGT		[155]
Thr126	CSD		Ser102 phosphorylation (↑)
Ser209, Ser313	CTD
Phosphorylation
Thr58, Thr67, Thr78 (MSY2)	CSD	CDK1	RNA or mRNP protein binding (↓)	[130]
Tyr99	CSD	Akt/PI3K pathway (?)	??	[156,157]
Ser99 (chk-YB-1) (Ser102 analog)	CSD	Akt	YB-1 (Ser99E, Ser99A) translation inhibition (↓) Nuclear translocation (n/e)	[158]
Ser102	CSD	Akt RSK (CN, calcineurin dephosphorylates)	Changes in YB-1-mRNA binding targets	[75,107]
			YB-1 binding to cap-adjusted mRNA regions (↓) Nuclear translocation (n/e)	[75]
			Nuclear translocation (↑)	[159,160,161]
			Promoter binding, incl. CCL5, EGFR, HER2 (↑?)	[159,162,163]
Ser134 (YB-3) (S102 analog)	CSD	Akt, RSK	??	[164]
Ser165	CTD	CKII (?)	Nuclear translocation (↑) Activation of NF-κB(↑?)	[165,166]
Ser176	CTD	CKI	Nuclear translocation (↑) Activation of NF-κB(↑?)	[167]
Thr188	CTD (NLS-2)	??	??	[168]
Thr281	CTD (NLS-3)	??	Nuclear translocation (↑?)	[168]
Poly(ADP-ribosyl)ation
??	CTD (219-324)	PARP1, PARP2	DNA binding (↓) APE1 activity (↑)	[169,170]
SUMOylation
Putative DSKA(287-290), TKED 60-63, EKRE (151-154) (zYB-1)	??	??	Nuclear translocation (↑?)	[171]
Ubiquitination
Lys27	A/P	HACE1	YB-1 secretion (↑)	[172]
Lys48	A/P	?? (OTUB1, deubiquitinase)	YB-1 protein stability (↓)	[173]