Ser9

Site Information
EEPQsDPsVEPPLsQ SwissProt Entrez-Gene
Blast this site against: NCBI SwissProt PDB
Site Group ID: 447538

In vivo Characterization
Methods used to characterize site in vivo:	2D analysis ( 20 , 35 ) , immunoprecipitation ( 28 ) , mass spectrometry ( 3 , 6 , 9 ) , modification-specific antibody ( 22 , 34 ) , mutation of modification site ( 2 , 5 , 21 , 22 , 25 , 30 ) , phospho-antibody ( 1 , 2 , 4 , 5 , 7 , 8 , 10 , 12 , 14 , 15 , 16 , 17 , 18 , 20 , 22 , 23 , 24 , 25 , 26 , 27 , 28 , 29 , 31 , 32 , 33 , 34 ) , phosphopeptide mapping ( 35 ) , western blotting ( 1 , 2 , 4 , 5 , 7 , 8 , 10 , 12 , 14 , 16 , 20 , 22 , 24 , 25 , 32 , 33 , 34 , 35 )
Disease tissue studied:	ataxia-telangiectasia ( 16 , 28 ) , bone cancer ( 5 , 20 ) , brain cancer ( 17 , 35 ) , glioblastoma ( 17 , 35 ) , glioblastoma multiforme ( 35 ) , glioma ( 17 , 35 ) , breast cancer ( 12 ) , colorectal cancer ( 14 , 21 , 22 ) , colorectal carcinoma ( 14 , 21 , 22 ) , leukemia ( 10 ) , acute lymphocytic leukemia ( 10 ) , liver cancer ( 4 ) , hepatocellular carcinoma ( 4 ) , lung cancer ( 1 , 2 , 5 , 6 , 12 , 14 , 22 , 24 , 33 , 35 ) , non-small cell lung cancer ( 1 , 2 , 5 , 6 , 12 , 14 , 22 , 35 ) , non-small cell lung adenocarcinoma ( 2 , 6 , 35 ) , non-small cell squamous cell lung carcinoma ( 14 ) , neuroblastoma ( 8 , 28 )
Relevant cell line - cell type - tissue:	293 (epithelial) [AT1 (human), transfection, AT1R stable transfected HEK293] ( 9 ) , A2182 ( 32 ) , A549 (pulmonary) ( 14 , 22 , 24 , 33 ) , Aspc1 (pancreatic) ( 32 ) , AT1ABR (lymphoblastoid) ( 16 ) , AT24RM (lymphoblastoid) ( 28 ) , AT2KY ( 29 ) , AT5BI (fibroblast) ( 29 ) , BT (epithelial) ( 16 , 28 ) , C3ABR (lymphoblastoid) ( 16 , 28 ) , Calu 6 (pulmonary) ( 35 ) , chondrocyte ( 25 ) , COLO-320 (intestinal) ( 32 ) , COS (fibroblast) ( 30 ) , Flp-In T-Rex-293 (epithelial) [PRKD1 (human), genetic knockin] ( 3 ) , GM00637 (lymphoblast) ( 31 ) , GM01526 (lymphoblast) ( 28 ) , GM02254 (lymphoblast) ( 28 ) , GM638 (fibroblast) ( 29 ) , HCC (hepatic) ( 4 ) , HCT116 (intestinal) ( 14 , 21 , 22 ) , HE49 (embryonic) ( 29 ) , hepatocyte-liver ( 4 ) , HepG2 (hepatic) ( 4 ) , IMR-90 (fibroblast) ( 26 ) , IMR32 (neural crest) ( 8 ) , L3 (lymphoblastoid) ( 28 ) , M059J (glial) ( 17 ) , MDA-MB-231 (breast cell) ( 12 , 23 ) , MEF (fibroblast) ( 22 ) , MRC5 (fibroblast) ( 22 ) , MT1 (lymphoblastoid) ( 18 ) , NALM6 (B lymphocyte) ( 10 ) , NCI-H1299 (pulmonary) ( 1 , 2 , 5 , 12 , 14 , 22 , 30 ) , NCI-H1650 (pulmonary) ( 14 ) , NCI-H1703 (squamous) ( 14 ) , NCI-H1975 (pulmonary) ( 2 ) , NCI-H23 (pulmonary) ( 14 ) , NCI-H460 (pulmonary) ( 14 ) , NCI-H596 (pulmonary) ( 32 ) , NCI-H838 (pulmonary) ( 6 ) , NHF (fibroblast) ( 34 ) , OM431 ( 32 ) , Saos-2 (bone cell) ( 5 ) , SHEP (neuron) ( 28 ) , SJSA-1 (bone cell) ( 32 ) , skin ( 27 ) , SW680 (intestinal) ( 32 ) , T98G (glial) [p53 (human), transfection] ( 35 ) , TK6 (lymphoblastoid) ( 18 ) , U2OS (bone cell) [GR (human)] ( 15 ) , U2OS (bone cell) ( 20 ) , WM4 ( 32 ) , WM5 ( 32 ) , WM793 ( 32 ) , WS1 (fibroblast) ( 22 )

Upstream Regulation
Regulatory protein:	DUSP26 (human) ( 8 ) , LKB1 (human) ( 14 )
Kinases, in vitro:	CK1D (human) ( 33 ) , DNAPK (human) ( 17 , 19 )
Treatments:	adriamycin ( 10 , 16 , 20 , 22 , 28 , 33 ) , asbestos ( 24 ) , colforsin ( 10 ) , Go_6976 ( 4 ) , H2O2 ( 31 ) , heat_shock ( 29 ) , IBMX ( 10 ) , ionizing_radiation ( 22 , 28 , 29 , 33 ) , morphine ( 23 ) , nocodazole ( 22 ) , PALA ( 22 ) , PD98059 ( 12 ) , siRNA ( 14 ) , SNP ( 25 ) , taxol ( 22 ) , TMZ ( 18 ) , UV ( 22 , 26 , 27 , 32 , 33 , 34 ) , virus infection ( 4 )

Downstream Regulation
Effects of modification on p53:	acetylation ( 22 , 28 ) , intracellular localization ( 1 ) , phosphorylation ( 22 )
Effects of modification on biological processes:	apoptosis, altered ( 14 , 18 ) , apoptosis, induced ( 21 ) , carcinogenesis, inhibited ( 1 ) , cell cycle regulation ( 18 ) , transcription, altered ( 21 )

References
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2	Vaughan CA, et al. (2021) The oncogenicity of tumor-derived mutant p53 is enhanced by the recruitment of PLK3. Nat Commun 12, 704 33514736 Curated Info
3	Franz-Wachtel M, et al. (2012) Global detection of protein kinase D-dependent phosphorylation events in nocodazole-treated human cells. Mol Cell Proteomics 11, 160-70 22496350 Curated Info
4	Knoll S, et al. (2011) Dissection of cell context-dependent interactions between HBx and p53 family members in regulation of apoptosis: a role for HBV-induced HCC. Cell Cycle 10, 3554-65 22030623 Curated Info
5	Ozeki C, et al. (2011) Cancer susceptibility polymorphism of p53 at codon 72 affects phosphorylation and degradation of p53 protein. J Biol Chem 286, 18251-60 21454683 Curated Info
6	Rikova K (2011) CST Curation Set: 10883; Year: 2011; Biosample/Treatment: cell line, NCI-H838/untreated; Disease: non-small cell lung cancer; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[STY]) Curated Info
7	Park J, et al. (2010) Dyrk1A phosphorylates p53 and inhibits proliferation of embryonic neuronal cells. J Biol Chem 285, 31895-906 20696760 Curated Info
8	Shang X, et al. (2010) Dual-specificity phosphatase 26 is a novel p53 phosphatase and inhibits p53 tumor suppressor functions in human neuroblastoma. Oncogene 29, 4938-46 20562916 Curated Info
9	Christensen GL, et al. (2010) Quantitative phosphoproteomics dissection of seven-transmembrane receptor signaling using full and biased agonists. Mol Cell Proteomics 9, 1540-53 20363803 Curated Info
10	Safa M, et al. (2010) Inhibitory role of cAMP on doxorubicin-induced apoptosis in pre-B ALL cells through dephosphorylation of p53 serine residues. Apoptosis 15, 196-203 19882354 Curated Info
11	Nishimura T, et al. (2009) Hepatitis C virus impairs p53 via persistent overexpression of 3beta-hydroxysterol Delta24-reductase. J Biol Chem 284, 36442-52 19861417 Curated Info
12	Adorno M, et al. (2009) A Mutant-p53/Smad complex opposes p63 to empower TGFbeta-induced metastasis. Cell 137, 87-98 19345189 Curated Info
13	Warnock LJ, et al. (2008) Influence of tetramerisation on site-specific post-translational modifications of p53: comparison of human and murine p53 tumor suppressor protein. Cancer Biol Ther 7, 1481-9 18769132 Curated Info
14	Zhong D, et al. (2008) LKB1 is necessary for Akt-mediated phosphorylation of proapoptotic proteins. Cancer Res 68, 7270-7 18794113 Curated Info
15	Mayo LD, et al. (2005) Phosphorylation of human p53 at serine 46 determines promoter selection and whether apoptosis is attenuated or amplified. J Biol Chem 280, 25953-9 15843377 Curated Info
16	Kurz EU, Douglas P, Lees-Miller SP (2004) Doxorubicin activates ATM-dependent phosphorylation of multiple downstream targets in part through the generation of reactive oxygen species. J Biol Chem 279, 53272-81 15489221 Curated Info
17	Komiyama S, et al. (2004) Potentiality of DNA-dependent protein kinase to phosphorylate Ser46 of human p53. Biochem Biophys Res Commun 323, 816-22 15381073 Curated Info
18	Caporali S, et al. (2004) DNA damage induced by temozolomide signals to both ATM and ATR: role of the mismatch repair system. Mol Pharmacol 66, 478-91 15322239 Curated Info
19	Soubeyrand S, Schild-Poulter C, Haché RJ (2004) Structured DNA promotes phosphorylation of p53 by DNA-dependent protein kinase at serine 9 and threonine 18. Eur J Biochem 271, 3776-84 15355354 Curated Info
20	Jackson MW, et al. (2004) Limited role of N-terminal phosphoserine residues in the activation of transcription by p53. Oncogene 23, 4477-87 15064747 Curated Info
21	Kaeser MD, Pebernard S, Iggo RD (2004) Regulation of p53 stability and function in HCT116 colon cancer cells. J Biol Chem 279, 7598-605 14665630 Curated Info
22	Saito S, et al. (2003) Phosphorylation site interdependence of human p53 post-translational modifications in response to stress. J Biol Chem 278, 37536-44 12860987 Curated Info
23	Tegeder I, et al. (2003) G protein-independent G1 cell cycle block and apoptosis with morphine in adenocarcinoma cells: involvement of p53 phosphorylation. Cancer Res 63, 1846-52 12702572 Curated Info
24	Matsuoka M, Igisu H, Morimoto Y (2003) Phosphorylation of p53 protein in A549 human pulmonary epithelial cells exposed to asbestos fibers. Environ Health Perspect 111, 509-12 12676607 Curated Info
25	Kim SJ, et al. (2002) p38 kinase regulates nitric oxide-induced apoptosis of articular chondrocytes by accumulating p53 via NFkappa B-dependent transcription and stabilization by serine 15 phosphorylation. J Biol Chem 277, 33501-8 12091386 Curated Info
26	Bulavin DV, et al. (2002) Amplification of PPM1D in human tumors abrogates p53 tumor-suppressor activity. Nat Genet 31, 210-5 12021785 Curated Info
27	Qin JZ, et al. (2002) Regulation of apoptosis by p53 in UV-irradiated human epidermis, psoriatic plaques and senescent keratinocytes. Oncogene 21, 2991-3002 12082529 Curated Info
28	Saito S, et al. (2002) ATM mediates phosphorylation at multiple p53 sites, including Ser(46), in response to ionizing radiation. J Biol Chem 277, 12491-4 11875057 Curated Info
29	Miyakoda M, Suzuki K, Kodama S, Watanabe M (2002) Activation of ATM and phosphorylation of p53 by heat shock. Oncogene 21, 1090-6 11850826 Curated Info
30	Dumaz N, Milne DM, Jardine LJ, Meek DW (2001) Critical roles for the serine 20, but not the serine 15, phosphorylation site and for the polyproline domain in regulating p53 turnover. Biochem J 359, 459-64 11583595 Curated Info
31	Xie S, et al. (2001) Reactive oxygen species-induced phosphorylation of p53 on serine 20 is mediated in part by polo-like kinase-3. J Biol Chem 276, 36194-9 11447225 Curated Info
32	Minamoto T, et al. (2001) Distinct pattern of p53 phosphorylation in human tumors. Oncogene 20, 3341-7 11423984 Curated Info
33	Higashimoto Y, et al. (2000) Human p53 is phosphorylated on serines 6 and 9 in response to DNA damage-inducing agents. J Biol Chem 275, 23199-203 10930428 Curated Info
34	Buschmann T, et al. (2000) p53 phosphorylation and association with murine double minute 2, c-Jun NH2-terminal kinase, p14ARF, and p300/CBP during the cell cycle and after exposure to ultraviolet irradiation. Cancer Res 60, 896-900 10706102 Curated Info
35	Ullrich SJ, et al. (1993) Phosphorylation at Ser-15 and Ser-392 in mutant p53 molecules from human tumors is altered compared to wild-type p53. Proc Natl Acad Sci U S A 90, 5954-8 8327466 Curated Info