Ser821

Site Information
GQLVGLNsPNsILkA SwissProt Entrez-Gene
Blast this site against: NCBI SwissProt PDB
Site Group ID: 467281

In vivo Characterization
Methods used to characterize site in vivo:	immunoprecipitation ( 2 , 4 , 12 , 15 , 30 ) , mass spectrometry ( 1 , 2 , 3 , 4 , 5 , 7 , 8 , 9 , 10 , 11 , 12 , 13 , 14 , 16 , 17 , 18 , 19 , 20 , 21 , 22 , 23 , 24 , 25 , 26 , 27 , 28 , 29 , 33 , 34 , 35 , 37 , 38 ) , mass spectrometry (in vitro) ( 31 ) , mutation of modification site ( 2 , 4 , 30 , 31 , 32 ) , phospho-antibody ( 2 , 4 , 12 , 15 , 30 ) , western blotting ( 2 , 4 , 12 , 15 )
Disease tissue studied:	bone cancer ( 2 ) , breast cancer ( 9 , 10 , 14 ) , breast ductal carcinoma ( 9 ) , HER2 positive breast cancer ( 5 ) , luminal A breast cancer ( 5 ) , luminal B breast cancer ( 5 ) , breast cancer, surrounding tissue ( 5 ) , breast cancer, triple negative ( 5 , 9 ) , cervical cancer ( 2 , 25 ) , cervical adenocarcinoma ( 2 , 25 ) , colorectal cancer ( 31 ) , colorectal carcinoma ( 31 ) , gastric cancer ( 23 ) , gastric carcinoma ( 23 ) , leukemia ( 16 ) , acute myelogenous leukemia ( 16 ) , lung cancer ( 14 , 18 ) , non-small cell lung cancer ( 14 , 18 ) , non-small cell large cell lung carcinoma ( 18 ) , ovarian cancer ( 9 ) , melanoma skin cancer ( 7 , 12 )
Relevant cell line - cell type - tissue:	293 (epithelial) ( 28 ) , breast ( 5 , 9 ) , BT-20 (breast cell) ( 14 ) , BT-549 (breast cell) ( 14 ) , H2009 (pulmonary) ( 14 ) , H2077 (pulmonary) ( 14 ) , H2887 (pulmonary) ( 14 ) , H322M (pulmonary) ( 14 ) , HCC1359 (pulmonary) ( 14 ) , HCC1937 (breast cell) ( 14 ) , HCC366 (pulmonary) ( 14 ) , HCC4006 (pulmonary) ( 14 ) , HCC78 (pulmonary) ( 14 ) , HCT116 (intestinal) ( 27 ) , HEK293T (epithelial) ( 2 , 12 , 15 ) , HeLa (cervical) ( 1 , 3 , 8 , 13 , 20 , 21 , 22 , 29 , 30 , 32 , 34 , 37 , 38 ) , HeLa S3 (cervical) ( 2 , 17 , 25 , 33 ) , HeLa_Meta (cervical) ( 24 ) , HeLa_Pro (cervical) ( 24 ) , HeLa_Telo (cervical) ( 24 ) , HMLER ('stem, breast cancer') [CXCR4 (human), knockdown] ( 10 ) , HMLER ('stem, breast cancer') ( 10 ) , HOP62 (pulmonary) ( 14 ) , HUES-7 ('stem, embryonic') ( 26 ) , HUES-9 ('stem, embryonic') ( 19 ) , Jurkat (T lymphocyte) ( 11 ) , K562 (erythroid) ( 13 , 35 ) , KG-1 (myeloid) ( 16 ) , MCF-7 (breast cell) ( 14 ) , MDA-MB-231 (breast cell) ( 14 ) , MDA-MB-435S (breast cell) ( 27 ) , MDA-MB-468 (breast cell) ( 14 ) , MEF (fibroblast) ( 4 ) , MKN-45 (gastric) ( 23 ) , MV4-11 (macrophage) ( 27 ) , NCI-H1395 (pulmonary) ( 14 ) , NCI-H157 (pulmonary) ( 14 ) , NCI-H1666 (pulmonary) ( 14 ) , NCI-H2030 (pulmonary) ( 14 ) , NCI-H322 (pulmonary) ( 14 ) , NCI-H520 (squamous) ( 14 ) , NCI-H661 (pulmonary) ( 18 ) , ovary ( 9 ) , PC9 (pulmonary) ( 14 ) , Sbcl2 (melanocyte) ( 12 ) , SW480 (intestinal) ( 31 ) , U2OS (bone cell) ( 2 ) , WM239A (melanocyte) ( 7 )

Upstream Regulation
Regulatory protein:	MAST1 (human) ( 4 )
Putative in vivo kinases:	ERK1 (human) ( 32 ) , ERK2 (human) ( 32 )
Kinases, in vitro:	TTK (human) ( 30 , 36 )
Phosphatases, in vitro:	PPP2R2A (human) ( 4 )
Treatments:	CXCL12 ( 10 ) , MG132_withdrawal ( 24 ) , nocodazole ( 4 , 25 , 33 ) , okadaic_acid ( 4 ) , thymidine ( 33 )

Downstream Regulation
Effects of modification on TTK:	intracellular localization ( 4 , 31 , 32 )

References
1	Hayward D, et al. (2019) CDK1-CCNB1 creates a spindle checkpoint-permissive state by enabling MPS1 kinetochore localization. J Cell Biol 30674582 Curated Info
2	Combes G, et al. (2018) Mps1 Phosphorylates Its N-Terminal Extension to Relieve Autoinhibition and Activate the Spindle Assembly Checkpoint. Curr Biol 28, 872-883.e5 29502948 Curated Info
3	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
4	Diril MK, et al. (2016) Loss of the Greatwall Kinase Weakens the Spindle Assembly Checkpoint. PLoS Genet 12, e1006310 27631493 Curated Info
5	Mertins P, et al. (2016) Proteogenomics connects somatic mutations to signalling in breast cancer. Nature 534, 55-62 27251275 Curated Info
6	Boeing S, et al. (2016) Multiomic Analysis of the UV-Induced DNA Damage Response. Cell Rep 15, 1597-1610 27184836 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	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
9	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
10	Yi T, et al. (2014) Quantitative phosphoproteomic analysis reveals system-wide signaling pathways downstream of SDF-1/CXCR4 in breast cancer stem cells. Proc Natl Acad Sci U S A 111, E2182-90 24782546 Curated Info
11	Mertins P, et al. (2013) Integrated proteomic analysis of post-translational modifications by serial enrichment. Nat Methods 10, 634-7 23749302 Curated Info
12	Liu J, et al. (2013) Phosphorylation of Mps1 by BRAFV600E prevents Mps1 degradation and contributes to chromosome instability in melanoma. Oncogene 32, 713-23 22430208 Curated Info
13	Zhou H, et al. (2013) Toward a comprehensive characterization of a human cancer cell phosphoproteome. J Proteome Res 12, 260-71 23186163 Curated Info
14	Klammer M, et al. (2012) Phosphosignature predicts dasatinib response in non-small cell lung cancer. Mol Cell Proteomics 11, 651-68 22617229 Curated Info
15	Lee S, et al. (2012) Characterization of spindle checkpoint kinase Mps1 reveals domain with functional and structural similarities to tetratricopeptide repeat motifs of Bub1 and BubR1 checkpoint kinases. J Biol Chem 287, 5988-6001 22187426 Curated Info
16	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
17	Santamaria A, et al. (2011) The Plk1-dependent phosphoproteome of the early mitotic spindle. Mol Cell Proteomics 10, M110.004457 20860994 Curated Info
18	Rikova K (2011) CST Curation Set: 10897; Year: 2011; Biosample/Treatment: cell line, NCI-H661/untreated; Disease: lung cancer; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[STY]) Curated Info
19	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
20	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
21	Hegemann B, et al. (2011) Systematic phosphorylation analysis of human mitotic protein complexes. Sci Signal 4, rs12 22067460 Curated Info
22	Zhou J (2010) CST Curation Set: 10708; Year: 2010; Biosample/Treatment: cell line, HeLa/untreated; Disease: cervical adenocarcinoma; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: PXpSP, pSPX(K/R) Antibodies Used to Purify Peptides prior to LCMS: Phospho-MAPK/CDK Substrates (PXSP or SPXR/K) (34B2) Rabbit mAb Cat#: 2325, PTMScan(R) Phospho-MAPK/CDK Substrate Motif (PXSP, SPXK/R) Immunoaffinity Beads Cat#: 1982 Curated Info
23	Moritz A (2010) CST Curation Set: 9979; Year: 2010; Biosample/Treatment: cell line, MKN-45/DMSO; Disease: gastric carcinoma; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[STY]) Curated Info
24	Dulla K, et al. (2010) Quantitative site-specific phosphorylation dynamics of human protein kinases during mitotic progression. Mol Cell Proteomics 9, 1167-81 20097925 Curated Info
25	Olsen JV, et al. (2010) Quantitative phosphoproteomics reveals widespread full phosphorylation site occupancy during mitosis. Sci Signal 3, ra3 20068231 Curated Info
26	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
27	Oppermann FS, et al. (2009) Large-scale proteomics analysis of the human kinome. Mol Cell Proteomics 8, 1751-64 19369195 Curated Info
28	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
29	Chen RQ, et al. (2009) CDC25B mediates rapamycin-induced oncogenic responses in cancer cells. Cancer Res 69, 2663-8 19276368 Curated Info
30	Tyler RK, et al. (2009) Phosphoregulation of human Mps1 kinase. Biochem J 417, 173-81 18680479 Curated Info
31	Xu Q, et al. (2009) Regulation of kinetochore recruitment of two essential mitotic spindle checkpoint proteins by Mps1 phosphorylation. Mol Biol Cell 20, 10-20 18923149 Curated Info
32	Borysova MK, Cui Y, Snyder M, Guadagno TM (2008) Knockdown of B-Raf impairs spindle formation and the mitotic checkpoint in human somatic cells. Cell Cycle 7, 2894-901 18787396 Curated Info
33	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
34	Dephoure N, et al. (2008) A quantitative atlas of mitotic phosphorylation. Proc Natl Acad Sci U S A 105, 10762-7 18669648 Curated Info
35	Stokes M (2008) CST Curation Set: 4605; 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
36	Jelluma N, et al. (2008) Chromosomal instability by inefficient Mps1 auto-activation due to a weakened mitotic checkpoint and lagging chromosomes. PLoS One 3, e2415 18545697 Curated Info
37	Kang J, Chen Y, Zhao Y, Yu H (2007) Autophosphorylation-dependent activation of human Mps1 is required for the spindle checkpoint. Proc Natl Acad Sci U S A 104, 20232-7 18083840 Curated Info
38	Yu LR, et al. (2007) Improved titanium dioxide enrichment of phosphopeptides from HeLa cells and high confident phosphopeptide identification by cross-validation of MS/MS and MS/MS/MS spectra. J Proteome Res 6, 4150-62 17924679 Curated Info