Ser321

Site Information
RLPLVPEsPRRMMTR SwissProt Entrez-Gene
Blast this site against: NCBI SwissProt PDB
Site Group ID: 470603

In vivo Characterization
Methods used to characterize site in vivo:	mass spectrometry ( 1 , 2 , 3 , 5 , 6 , 7 , 8 , 9 , 10 , 11 , 12 , 13 , 14 , 15 , 16 , 17 , 19 , 20 , 21 , 22 , 23 , 25 , 26 , 27 , 28 , 29 , 30 , 31 , 32 , 33 , 34 , 35 , 36 , 37 , 38 , 39 , 40 , 41 , 42 , 43 , 44 , 45 , 46 , 47 , 48 , 49 , 50 , 51 , 52 )
Disease tissue studied:	breast cancer ( 10 , 22 ) , breast ductal carcinoma ( 10 ) , HER2 positive breast cancer ( 3 ) , luminal A breast cancer ( 3 ) , luminal B breast cancer ( 3 ) , breast cancer, surrounding tissue ( 3 ) , breast cancer, triple negative ( 3 , 10 ) , cervical cancer ( 38 ) , cervical adenocarcinoma ( 38 ) , leukemia ( 25 , 49 ) , acute myelogenous leukemia ( 25 ) , acute erythroid leukemias, including erythroleukemia (M6a) and very rare pure erythroid leukemia (M6b) ( 21 ) , acute megakaryoblastic leukemia (M7) ( 21 ) , acute monoblastic leukemia (M5a) or acute monocytic leukemia (M5b) ( 21 ) , acute myeloblastic leukemia, with granulocytic maturation (M2) ( 21 ) , acute myeloblastic leukemia, without maturation (M1) ( 21 ) , chronic myelogenous leukemia ( 49 ) , lung cancer ( 8 , 15 , 22 , 32 , 46 ) , non-small cell lung cancer ( 22 , 46 ) , non-small cell lung adenocarcinoma ( 8 , 15 ) , lymphoma ( 11 ) , B cell lymphoma ( 21 ) , Burkitt's lymphoma ( 11 ) , non-Hodgkin's lymphoma ( 21 ) , follicular lymphoma ( 11 ) , mantle cell lymphoma ( 11 ) , neuroblastoma ( 20 ) , ovarian cancer ( 10 ) , pancreatic ductal adenocarcinoma ( 14 ) , multiple myeloma ( 21 ) , melanoma skin cancer ( 7 )
Relevant cell line - cell type - tissue:	'pancreatic, ductal'-pancreas ( 14 ) , 'stem, embryonic' ( 41 ) , 293 (epithelial) [AT1 (human), transfection, AT1R stable transfected HEK293] ( 36 ) , 293 (epithelial) [AT1 (human), transfection] ( 33 ) , 293 (epithelial) ( 42 ) , 293E (epithelial) ( 27 ) , 786-O (renal) [VHL (human), transfection] ( 5 ) , 786-O (renal) ( 5 ) , A549 (pulmonary) ( 16 ) , AML-193 (monocyte) ( 21 ) , BJAB (B lymphocyte) ( 11 ) , breast ( 3 , 10 ) , BT-20 (breast cell) ( 22 ) , BT-549 (breast cell) ( 22 ) , Calu 6 (pulmonary) ( 22 ) , CL1-0 (pulmonary) ( 32 ) , CL1-1 (pulmonary) ( 32 ) , CL1-2 (pulmonary) ( 32 ) , CL1-5 (pulmonary) ( 32 ) , CMK (megakaryoblast) ( 21 ) , CTS (myeloid) ( 21 ) , DOHH2 ('B lymphocyte, precursor') ( 21 ) , FL-18 (B lymphocyte) ( 11 ) , FL-318 (B lymphocyte) ( 11 ) , Flp-In T-Rex-293 (epithelial) [PRKD1 (human), genetic knockin] ( 23 ) , Flp-In T-Rex-293 (epithelial) ( 23 ) , H2009 (pulmonary) ( 22 ) , H2077 (pulmonary) ( 22 ) , H2887 (pulmonary) ( 22 ) , H322M (pulmonary) ( 22 ) , HCC1359 (pulmonary) ( 22 ) , HCC1937 (breast cell) ( 22 ) , HCC366 (pulmonary) ( 22 ) , HCC4006 (pulmonary) ( 22 ) , HCC78 (pulmonary) ( 22 ) , HEK293T (epithelial) ( 6 ) , HEL (erythroid) ( 21 ) , HeLa (cervical) ( 2 , 9 , 19 , 31 , 37 , 39 , 43 , 44 , 48 , 50 , 51 , 52 ) , HeLa S3 (cervical) ( 29 , 38 , 47 ) , HOP62 (pulmonary) ( 22 ) , HUES-7 ('stem, embryonic') ( 40 ) , HUES-9 ('stem, embryonic') ( 30 ) , JEKO-1 (B lymphocyte) ( 11 ) , Jurkat (T lymphocyte) ( 17 , 26 , 28 , 34 , 35 , 45 ) , K562 (erythroid) ( 19 , 39 , 49 ) , Kasumi-1 (myeloid) ( 21 ) , KG-1 (myeloid) ( 21 , 25 ) , liver ( 13 ) , lung ( 15 ) , MCF-7 (breast cell) ( 22 ) , MDA-MB-231 (breast cell) ( 22 ) , MDA-MB-468 (breast cell) ( 22 ) , MV4-11 (macrophage) ( 21 ) , NB10 (neural crest) ( 20 ) , NCEB-1 (B lymphocyte) ( 11 ) , NCI-H1299 (pulmonary) ( 46 ) , NCI-H1395 (pulmonary) ( 22 ) , NCI-H1568 (pulmonary) ( 22 ) , NCI-H157 (pulmonary) ( 22 ) , NCI-H1648 (pulmonary) ( 22 ) , NCI-H1666 (pulmonary) ( 22 ) , NCI-H2030 (pulmonary) ( 22 ) , NCI-H322 (pulmonary) ( 22 ) , NCI-H520 (squamous) ( 22 ) , NCI-H647 (pulmonary) ( 22 ) , NPC (neural crest) ( 20 ) , OCI-ly1 (B lymphocyte) ( 11 ) , OPM-2 (plasma cell) ( 21 ) , ovary ( 10 ) , P31/FUJ (erythroid) ( 21 ) , PC9 (pulmonary) ( 8 , 22 ) , PC9-IR (pulmonary) ( 8 ) , Raji (B lymphocyte) ( 11 ) , RAMOS (B lymphocyte) ( 11 ) , REC-1 (B lymphocyte) ( 11 ) , RL ('B lymphocyte, precursor') ( 21 ) , RPMI-8266 (plasma cell) ( 21 ) , SH-SY5Y (neural crest) [LRRK2 (human), transfection, over-expression of LRRK2(G2019S)] ( 12 ) , SH-SY5Y (neural crest) ( 12 ) , SU-DHL-4 (B lymphocyte) ( 11 ) , SU-DHL-6 (B lymphocyte) ( 21 ) , U266 (plasma cell) ( 21 ) , UPN-1 (B lymphocyte) ( 11 ) , Vero E6-S ('epithelial, kidney') ( 1 ) , WM239A (melanocyte) ( 7 )

Upstream Regulation
Treatments:	dasatinib ( 39 ) , ischemia ( 10 ) , nocodazole ( 38 )

References
1	Bouhaddou M, et al. (2020) The Global Phosphorylation Landscape of SARS-CoV-2 Infection. Cell 182 32645325 Curated Info
2	Huang H, et al. (2016) Simultaneous Enrichment of Cysteine-containing Peptides and Phosphopeptides Using a Cysteine-specific Phosphonate Adaptable Tag (CysPAT) in Combination with titanium dioxide (TiO2) Chromatography. Mol Cell Proteomics 15, 3282-3296 27281782 Curated Info
3	Mertins P, et al. (2016) Proteogenomics connects somatic mutations to signalling in breast cancer. Nature 534, 55-62 27251275 Curated Info
4	Boeing S, et al. (2016) Multiomic Analysis of the UV-Induced DNA Damage Response. Cell Rep 15, 1597-1610 27184836 Curated Info
5	Malec V, Coulson JM, Urbé S, Clague MJ (2015) Combined Analyses of the VHL and Hypoxia Signaling Axes in an Isogenic Pairing of Renal Clear Cell Carcinoma Cells. J Proteome Res 14, 5263-72 26506913 Curated Info
6	Franchin C, et al. (2015) Quantitative analysis of a phosphoproteome readily altered by the protein kinase CK2 inhibitor quinalizarin in HEK-293T cells. Biochim Biophys Acta 1854, 609-23 25278378 Curated Info
7	Stuart SA, et al. (2015) A Phosphoproteomic Comparison of B-RAFV600E and MKK1/2 Inhibitors in Melanoma Cells. Mol Cell Proteomics 14, 1599-615 25850435 Curated Info
8	Tsai CF, et al. (2015) Large-scale determination of absolute phosphorylation stoichiometries in human cells by motif-targeting quantitative proteomics. Nat Commun 6, 6622 25814448 Curated Info
9	Sharma K, et al. (2014) Ultradeep human phosphoproteome reveals a distinct regulatory nature of Tyr and Ser/Thr-based signaling. Cell Rep 8, 1583-94 25159151 Curated Info
10	Mertins P, et al. (2014) Ischemia in tumors induces early and sustained phosphorylation changes in stress kinase pathways but does not affect global protein levels. Mol Cell Proteomics 13, 1690-704 24719451 Curated Info
11	Rolland D, et al. (2014) Global phosphoproteomic profiling reveals distinct signatures in B-cell non-Hodgkin lymphomas. Am J Pathol 184, 1331-42 24667141 Curated Info
12	Luerman GC, et al. (2014) Phosphoproteomic evaluation of pharmacological inhibition of leucine-rich repeat kinase 2 reveals significant off-target effects of LRRK-2-IN-1. J Neurochem 128, 561-76 24117733 Curated Info
13	Bian Y, et al. (2014) An enzyme assisted RP-RPLC approach for in-depth analysis of human liver phosphoproteome. J Proteomics 96, 253-62 24275569 Curated Info
14	Britton D, et al. (2014) Quantification of pancreatic cancer proteome and phosphorylome: indicates molecular events likely contributing to cancer and activity of drug targets. PLoS One 9, e90948 24670416 Curated Info
15	Schweppe DK, Rigas JR, Gerber SA (2013) Quantitative phosphoproteomic profiling of human non-small cell lung cancer tumors. J Proteomics 91, 286-96 23911959 Curated Info
16	Kim JY, et al. (2013) Dissection of TBK1 signaling via phosphoproteomics in lung cancer cells. Proc Natl Acad Sci U S A 110, 12414-9 23836654 Curated Info
17	Mertins P, et al. (2013) Integrated proteomic analysis of post-translational modifications by serial enrichment. Nat Methods 10, 634-7 23749302 Curated Info
18	Shiromizu T, et al. (2013) Identification of missing proteins in the neXtProt database and unregistered phosphopeptides in the PhosphoSitePlus database as part of the Chromosome-centric Human Proteome Project. J Proteome Res 12, 2414-21 23312004 Curated Info
19	Zhou H, et al. (2013) Toward a comprehensive characterization of a human cancer cell phosphoproteome. J Proteome Res 12, 260-71 23186163 Curated Info
20	DeNardo BD, et al. (2013) Quantitative phosphoproteomic analysis identifies activation of the RET and IGF-1R/IR signaling pathways in neuroblastoma. PLoS One 8, e82513 24349301 Curated Info
21	Casado P, et al. (2013) Phosphoproteomics data classify hematological cancer cell lines according to tumor type and sensitivity to kinase inhibitors. Genome Biol 14, R37 23628362 Curated Info
22	Klammer M, et al. (2012) Phosphosignature predicts dasatinib response in non-small cell lung cancer. Mol Cell Proteomics 11, 651-68 22617229 Curated Info
23	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
24	Beli P, et al. (2012) Proteomic Investigations Reveal a Role for RNA Processing Factor THRAP3 in the DNA Damage Response. Mol Cell 46, 212-25 22424773 Curated Info
25	Weber C, Schreiber TB, Daub H (2012) Dual phosphoproteomics and chemical proteomics analysis of erlotinib and gefitinib interference in acute myeloid leukemia cells. J Proteomics 75, 1343-56 22115753 Curated Info
26	Mulhern D (2011) CST Curation Set: 12713; Year: 2011; Biosample/Treatment: cell line, Jurkat/calyculin_A & pervanadate; Disease: T cell leukemia; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[ST]XP Antibodies Used to Purify Peptides prior to LCMS: Phospho-(Ser) 14-3-3 Binding Motif Antibody Cat#: 9601 Curated Info
27	Hsu PP, et al. (2011) The mTOR-regulated phosphoproteome reveals a mechanism of mTORC1-mediated inhibition of growth factor signaling. Science 332, 1317-22 21659604 Curated Info
28	Guo A (2011) CST Curation Set: 11989; Year: 2011; Biosample/Treatment: cell line, Jurkat/calyculin_A & pervanadate; Disease: T cell leukemia; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[ST]XP Antibodies Used to Purify Peptides prior to LCMS: Phospho-(Ser) 14-3-3 Binding Motif Antibody Cat#: 9601 Curated Info
29	Santamaria A, et al. (2011) The Plk1-dependent phosphoproteome of the early mitotic spindle. Mol Cell Proteomics 10, M110.004457 20860994 Curated Info
30	Rigbolt KT, et al. (2011) System-wide temporal characterization of the proteome and phosphoproteome of human embryonic stem cell differentiation. Sci Signal 4, rs3 21406692 Curated Info
31	Kettenbach AN, et al. (2011) Quantitative phosphoproteomics identifies substrates and functional modules of aurora and polo-like kinase activities in mitotic cells. Sci Signal 4, rs5 21712546 Curated Info
32	Wang YT, et al. (2010) An informatics-assisted label-free quantitation strategy that depicts phosphoproteomic profiles in lung cancer cell invasion. J Proteome Res 9, 5582-97 20815410 Curated Info
33	Xiao K, et al. (2010) Global phosphorylation analysis of beta-arrestin-mediated signaling downstream of a seven transmembrane receptor (7TMR). Proc Natl Acad Sci U S A 107, 15299-304 20686112 Curated Info
34	Possemato A (2010) CST Curation Set: 10111; Year: 2010; Biosample/Treatment: cell line, Jurkat/calyculin_A & pervanadate; Disease: T cell leukemia; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: RXXp[ST] Antibodies Used to Purify Peptides prior to LCMS: Phospho-Akt Substrate (RXRXXS/T) (110B7) Rabbit mAb Cat#: 9614, PTMScan(R) Phospho-Akt Substrate Motif (RXXS/T) Immunoaffinity Beads Cat#: 1978 Curated Info
35	Possemato A (2010) CST Curation Set: 10110; Year: 2010; Biosample/Treatment: cell line, Jurkat/calyculin_A & pervanadate; Disease: T cell leukemia; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: RXXp[ST] Antibodies Used to Purify Peptides prior to LCMS: Phospho-Akt Substrate (RXRXXS/T) (110B7) Rabbit mAb Cat#: 9614, PTMScan(R) Phospho-Akt Substrate Motif (RXXS/T) Immunoaffinity Beads Cat#: 1978 Curated Info
36	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
37	Moritz A (2010) CST Curation Set: 9243; Year: 2010; Biosample/Treatment: cell line, HeLa/UV; Disease: cervical adenocarcinoma; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[STY]) Curated Info
38	Olsen JV, et al. (2010) Quantitative phosphoproteomics reveals widespread full phosphorylation site occupancy during mitosis. Sci Signal 3, ra3 20068231 Curated Info
39	Pan C, Olsen JV, Daub H, Mann M (2009) Global effects of kinase inhibitors on signaling networks revealed by quantitative phosphoproteomics. Mol Cell Proteomics 8, 2796-808 19651622 Curated Info
40	Van Hoof D, et al. (2009) Phosphorylation dynamics during early differentiation of human embryonic stem cells. Cell Stem Cell 5, 214-26 19664995 Curated Info
41	Brill LM, et al. (2009) Phosphoproteomic analysis of human embryonic stem cells. Cell Stem Cell 5, 204-13 19664994 Curated Info
42	Gauci S, et al. (2009) Lys-N and trypsin cover complementary parts of the phosphoproteome in a refined SCX-based approach. Anal Chem 81, 4493-501 19413330 Curated Info
43	Chen Y, et al. (2009) Combined integrin phosphoproteomic analyses and small interfering RNA--based functional screening identify key regulators for cancer cell adhesion and migration. Cancer Res 69, 3713-20 19351860 Curated Info
44	Chen RQ, et al. (2009) CDC25B mediates rapamycin-induced oncogenic responses in cancer cells. Cancer Res 69, 2663-8 19276368 Curated Info
45	Mayya V, et al. (2009) Quantitative phosphoproteomic analysis of T cell receptor signaling reveals system-wide modulation of protein-protein interactions. Sci Signal 2, ra46 19690332 Curated Info
46	Tsai CF, et al. (2008) Immobilized metal affinity chromatography revisited: pH/acid control toward high selectivity in phosphoproteomics. J Proteome Res 7, 4058-69 18707149 Curated Info
47	Daub H, et al. (2008) Kinase-selective enrichment enables quantitative phosphoproteomics of the kinome across the cell cycle. Mol Cell 31, 438-48 18691976 Curated Info
48	Dephoure N, et al. (2008) A quantitative atlas of mitotic phosphorylation. Proc Natl Acad Sci U S A 105, 10762-7 18669648 Curated Info
49	Stokes M (2008) CST Curation Set: 4391; Year: 2008; Biosample/Treatment: cell line, K562/untreated; Disease: chronic myelogenous leukemia; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[STY]) Curated Info
50	Cantin GT, et al. (2008) Combining protein-based IMAC, peptide-based IMAC, and MudPIT for efficient phosphoproteomic analysis. J Proteome Res 7, 1346-51 18220336 Curated Info
51	Olsen JV, et al. (2006) Global, in vivo, and site-specific phosphorylation dynamics in signaling networks. Cell 127, 635-48 17081983 Curated Info
52	Beausoleil SA, et al. (2006) A probability-based approach for high-throughput protein phosphorylation analysis and site localization. Nat Biotechnol 24, 1285-92 16964243 Curated Info