Thr514

Site Information
sVtPktVtPAssAkt SwissProt Entrez-Gene
Blast this site against: NCBI SwissProt PDB
Site Group ID: 454386

In vivo Characterization
Methods used to characterize site in vivo:	mass spectrometry ( 3 , 7 , 9 , 10 , 11 , 13 , 14 , 15 , 16 , 17 , 18 , 19 , 20 , 21 , 23 , 24 , 25 , 26 , 27 , 29 , 30 , 31 , 32 , 33 , 34 , 35 , 36 , 37 , 38 , 39 , 40 , 41 , 42 , 43 , 44 , 45 , 46 , 48 , 49 , 50 , 52 ) , mutation of modification site ( 6 , 28 , 51 ) , phospho-antibody ( 1 , 5 , 6 , 12 , 22 , 47 , 51 ) , western blotting ( 1 , 5 , 6 , 12 , 22 , 47 , 51 )
Disease tissue studied:	bipolar disprder ( 1 ) , brain cancer ( 13 , 14 , 15 , 16 , 17 , 18 , 19 , 20 , 21 ) , neuroblastoma ( 25 , 51 ) , neuroendocrine cancer ( 13 , 14 , 15 , 16 , 17 , 18 , 19 , 20 , 21 )
Relevant cell line - cell type - tissue:	'3T3-L1, differentiated' (adipocyte) ( 7 ) , 'brain, cerebral cortex' ( 29 , 47 ) , 'brain, embryonic' ( 23 , 30 , 32 , 35 , 36 , 39 , 40 , 42 , 43 , 46 , 48 ) , 'brain, forebrain' ( 45 ) , 'brain, hippocampus' ( 1 , 47 , 51 ) , 'fat, brown' ( 26 ) , 'neuron, cortical' ( 28 ) , 'stem, induced pluripotent' ( 1 ) , brain ( 10 , 13 , 14 , 15 , 16 , 17 , 18 , 19 , 20 , 21 , 26 , 27 , 31 , 33 , 34 , 37 , 38 , 41 , 44 , 49 , 50 , 52 ) , COS (fibroblast) ( 6 ) , heart ( 26 ) , liver ( 26 ) , lung ( 26 ) , MEF (fibroblast) ( 9 ) , MEF (fibroblast) [TSC2 (mouse), homozygous knockout] ( 24 ) , N1E-115 (neuron) ( 25 , 51 ) , neuron-'brain, cerebral cortex' ( 5 , 51 ) , neuron-'brain, hippocampus' ( 22 ) , neuron-dorsal root ganglia ( 6 , 12 ) , pancreas ( 26 ) , PC-12 (chromaffin) [TrkA (rat), transfection] ( 11 ) , RAW 264.7 (macrophage) ( 3 ) , SH-SY5Y (neural crest) ( 51 ) , spleen ( 26 ) , testis ( 26 )

Upstream Regulation
Regulatory protein:	APP (human) ( 47 ) , APP (mouse) ( 47 ) , axin 1 (mouse) ( 22 ) , CDK5 (human) ( 47 ) , CDK5 (mouse) ( 51 ) , PSEN1 (mouse) ( 47 )
Putative in vivo kinases:	GSK3A (human) ( 5 ) , GSK3B (mouse) ( 51 )
Treatments:	bisindolylmaleimide ( 6 ) , CT-99021 ( 51 ) , insulin ( 7 ) , kainic_acid ( 6 ) , lithium ( 1 ) , LPA ( 25 ) , LY294002 ( 7 ) , metamfetamine ( 1 ) , MG132 ( 12 ) , MK-2206 ( 7 ) , nerve_damage ( 12 ) , Sem3A ( 51 ) , TDZD-8 ( 12 )

Downstream Regulation
Effects of modification on biological processes:	cell growth, altered ( 28 ) , cell motility, induced ( 5 ) , cytoskeletal reorganization ( 5 )

References
1	Tobe BTD, et al. (2017) Probing the lithium-response pathway in hiPSCs implicates the phosphoregulatory set-point for a cytoskeletal modulator in bipolar pathogenesis. Proc Natl Acad Sci U S A 114, E4462-E4471 28500272 Curated Info
2	Sacco F, et al. (2016) Glucose-regulated and drug-perturbed phosphoproteome reveals molecular mechanisms controlling insulin secretion. Nat Commun 7, 13250 27841257 Curated Info
3	Pinto SM, et al. (2015) Quantitative phosphoproteomic analysis of IL-33-mediated signaling. Proteomics 15, 532-44 25367039 Curated Info
4	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
5	Morgan-Smith M, et al. (2014) GSK-3 signaling in developing cortical neurons is essential for radial migration and dendritic orientation. Elife 3, e02663 25073924 Curated Info
6	Marques JM, et al. (2013) CRMP2 Tethers Kainate Receptor Activity to Cytoskeleton Dynamics during Neuronal Maturation. J Neurosci 33, 18298-310 24227739 Curated Info
7	Humphrey SJ, et al. (2013) Dynamic Adipocyte Phosphoproteome Reveals that Akt Directly Regulates mTORC2. Cell Metab 17, 1009-20 23684622 Curated Info
8	Qin Z, et al. (2013) Isoaspartate accumulation in mouse brain is associated with altered patterns of protein phosphorylation and acetylation, some of which are highly sex-dependent. PLoS One 8, e80758 24224061 Curated Info
9	Wu X, et al. (2012) Investigation of receptor interacting protein (RIP3)-dependent protein phosphorylation by quantitative phosphoproteomics. Mol Cell Proteomics 11, 1640-51 22942356 Curated Info
10	Goswami T, et al. (2012) Comparative phosphoproteomic analysis of neonatal and adult murine brain. Proteomics 12, 2185-9 22807455 Curated Info
11	Biarc J, Chalkley RJ, Burlingame AL, Bradshaw RA (2012) The induction of serine/threonine protein phosphorylations by a PDGFR/TrkA chimera in stably transfected PC12 cells. Mol Cell Proteomics 11, 15-30 22027198 Curated Info
12	Wakatsuki S, Saitoh F, Araki T (2011) ZNRF1 promotes Wallerian degeneration by degrading AKT to induce GSK3B-dependent CRMP2 phosphorylation. Nat Cell Biol 13, 1415-23 22057101 Curated Info
13	Guo A (2011) CST Curation Set: 12728; Year: 2011; Biosample/Treatment: tissue, brain/untreated; Disease: brain cancer; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[ST]P Curated Info
14	Guo A (2011) CST Curation Set: 12729; Year: 2011; Biosample/Treatment: tissue, brain/untreated; Disease: brain cancer; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[ST]P Curated Info
15	Guo A (2011) CST Curation Set: 12730; Year: 2011; Biosample/Treatment: tissue, brain/untreated; Disease: brain cancer; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[ST]P Curated Info
16	Guo A (2011) CST Curation Set: 12732; Year: 2011; Biosample/Treatment: tissue, brain/untreated; Disease: brain cancer; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[ST]P Antibodies Used to Purify Peptides prior to LCMS: Phospho-(Ser) CDKs Substrate Antibody Cat#: 2324, PTMScan(R) Phospho-CDK Substrate Motif (K/RS*PXK/R) Immunoaffinity Beads Cat#: 1981 Curated Info
17	Guo A (2011) CST Curation Set: 12733; Year: 2011; Biosample/Treatment: tissue, brain/untreated; Disease: brain cancer; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[ST]P Antibodies Used to Purify Peptides prior to LCMS: Phospho-(Ser) CDKs Substrate Antibody Cat#: 2324, PTMScan(R) Phospho-CDK Substrate Motif (K/RS*PXK/R) Immunoaffinity Beads Cat#: 1981 Curated Info
18	Guo A (2011) CST Curation Set: 12734; Year: 2011; Biosample/Treatment: tissue, brain/untreated; Disease: brain cancer; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[ST]P Antibodies Used to Purify Peptides prior to LCMS: Phospho-(Ser) CDKs Substrate Antibody Cat#: 2324, PTMScan(R) Phospho-CDK Substrate Motif (K/RS*PXK/R) Immunoaffinity Beads Cat#: 1981 Curated Info
19	Guo A (2011) CST Curation Set: 12735; Year: 2011; Biosample/Treatment: tissue, brain/untreated; Disease: brain cancer; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[ST]P Antibodies Used to Purify Peptides prior to LCMS: Phospho-(Ser) CDKs Substrate Antibody Cat#: 2324, PTMScan(R) Phospho-CDK Substrate Motif (K/RS*PXK/R) Immunoaffinity Beads Cat#: 1981 Curated Info
20	Guo A (2011) CST Curation Set: 12738; Year: 2011; Biosample/Treatment: tissue, brain/untreated; Disease: brain cancer; 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
21	Guo A (2011) CST Curation Set: 12739; Year: 2011; Biosample/Treatment: tissue, brain/untreated; Disease: brain cancer; 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
22	Fang WQ, et al. (2011) Cdk5-mediated phosphorylation of Axin directs axon formation during cerebral cortex development. J Neurosci 31, 13613-24 21940452 Curated Info
23	Guo A (2011) CST Curation Set: 12476; Year: 2011; Biosample/Treatment: tissue, brain/untreated; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[STY] Curated Info
24	Yu Y, et al. (2011) Phosphoproteomic analysis identifies Grb10 as an mTORC1 substrate that negatively regulates insulin signaling. Science 332, 1322-6 21659605 Curated Info
25	Wang Y, et al. (2011) Spatial phosphoprotein profiling reveals a compartmentalized extracellular signal-regulated kinase switch governing neurite growth and retraction. J Biol Chem 286, 18190-201 21454597 Curated Info
26	Huttlin EL, et al. (2010) A tissue-specific atlas of mouse protein phosphorylation and expression. Cell 143, 1174-89 21183079 Curated Info
27	Wiśniewski JR, et al. (2010) Brain phosphoproteome obtained by a FASP-based method reveals plasma membrane protein topology. J Proteome Res 9, 3280-9 20415495 Curated Info
28	Shinkai-Ouchi F, et al. (2010) Identification and structural analysis of C-terminally truncated collapsin response mediator protein-2 in a murine model of prion diseases. Proteome Sci 8, 53 20961402 Curated Info
29	Tweedie-Cullen RY, Reck JM, Mansuy IM (2009) Comprehensive mapping of post-translational modifications on synaptic, nuclear, and histone proteins in the adult mouse brain. J Proteome Res 8, 4966-82 19737024 Curated Info
30	Zhou J (2009) CST Curation Set: 7441; Year: 2009; Biosample/Treatment: tissue, brain/untreated; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[ST]P Curated Info
31	Zhou J (2009) CST Curation Set: 7443; Year: 2009; Biosample/Treatment: tissue, brain/untreated; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[ST]P Curated Info
32	Zhou J (2009) CST Curation Set: 7440; Year: 2009; Biosample/Treatment: tissue, brain/untreated; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[ST]P Curated Info
33	Zhou J (2009) CST Curation Set: 7442; Year: 2009; Biosample/Treatment: tissue, brain/untreated; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[ST]P Curated Info
34	Zhou J (2009) CST Curation Set: 7431; Year: 2009; Biosample/Treatment: tissue, brain/untreated; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: pSP Curated Info
35	Zhou J (2009) CST Curation Set: 7428; Year: 2009; Biosample/Treatment: tissue, brain/untreated; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: pSP Curated Info
36	Zhou J (2009) CST Curation Set: 7429; Year: 2009; Biosample/Treatment: tissue, brain/untreated; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: pSP Curated Info
37	Zhou J (2009) CST Curation Set: 7413; Year: 2009; Biosample/Treatment: tissue, brain/untreated; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: HXXp[ST] Curated Info
38	Zhou J (2009) CST Curation Set: 7414; Year: 2009; Biosample/Treatment: tissue, brain/untreated; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: HXXp[ST] Curated Info
39	Zhou J (2009) CST Curation Set: 7412; Year: 2009; Biosample/Treatment: tissue, brain/untreated; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: HXXp[ST] Curated Info
40	Zhou J (2009) CST Curation Set: 7411; Year: 2009; Biosample/Treatment: tissue, brain/untreated; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: HXXp[ST] Curated Info
41	Zhou J (2008) CST Curation Set: 4863; Year: 2008; Biosample/Treatment: tissue, brain/untreated; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[ST]P Antibodies Used to Purify Peptides prior to LCMS: Phospho-Thr-Pro Motif Antibody (polyAB) Cat#: 3003, PTMScan(R) Phospho-Thr-Pro Motif (T*P) Immunoaffinity Beads Cat#: 1996 Curated Info
42	Zhou J (2008) CST Curation Set: 4864; Year: 2008; Biosample/Treatment: tissue, brain/untreated; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[ST]P Antibodies Used to Purify Peptides prior to LCMS: Phospho-Thr-Pro Motif Antibody (polyAB) Cat#: 3003, PTMScan(R) Phospho-Thr-Pro Motif (T*P) Immunoaffinity Beads Cat#: 1996 Curated Info
43	Zhou J (2008) CST Curation Set: 4860; Year: 2008; Biosample/Treatment: tissue, brain/untreated; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[ST]P Curated Info
44	Zhou J (2008) CST Curation Set: 4779; Year: 2008; Biosample/Treatment: tissue, brain/untreated; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: pTXR Antibodies Used to Purify Peptides prior to LCMS: Phospho-Threonine-X-Arginine Antibody Cat#: 2351, PTMScan(R) Phospho-Thr-X-Arg Motif (T*XR) Immunoaffinity Beads Cat#: 1988 Curated Info
45	Collins MO, et al. (2008) Phosphoproteomic analysis of the mouse brain cytosol reveals a predominance of protein phosphorylation in regions of intrinsic sequence disorder. Mol Cell Proteomics 7, 1331-48 18388127 Curated Info
46	Zhou J (2008) CST Curation Set: 4323; Year: 2008; Biosample/Treatment: tissue, brain/untreated; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: pY Curated Info
47	Cole AR, et al. (2007) Collapsin response mediator protein-2 hyperphosphorylation is an early event in Alzheimer's disease progression. J Neurochem 103, 1132-44 17683481 Curated Info
48	Guo A (2007) CST Curation Set: 2715; Year: 2007; Biosample/Treatment: tissue, brain/-; Disease: -; 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
49	Guo A (2007) CST Curation Set: 2718; Year: 2007; Biosample/Treatment: tissue, brain/-; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[ST](D/E)X(D/E) Curated Info
50	Guo A (2007) CST Curation Set: 2682; Year: 2007; Biosample/Treatment: tissue, brain/-; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[ST]Q Antibodies Used to Purify Peptides prior to LCMS: Phospho-(Ser/Thr) ATM/ATR Substrate Antibody Cat#: 2851 Curated Info
51	Cole AR, et al. (2006) Distinct priming kinases contribute to differential regulation of collapsin response mediator proteins by glycogen synthase kinase-3 in vivo. J Biol Chem 281, 16591-8 16611631 Curated Info
52	Trinidad JC, et al. (2006) Comprehensive identification of phosphorylation sites in postsynaptic density preparations. Mol Cell Proteomics 5, 914-22 16452087 Curated Info