Ser86

Site Information
RALNRQLssGVsEIR SwissProt Entrez-Gene
Blast this site against: NCBI SwissProt PDB
Site Group ID: 448403

In vivo Characterization
Methods used to characterize site in vivo:	2D analysis ( 46 ) , [32P] ATP in vitro ( 46 ) , [32P] bio-synthetic labeling ( 42 ) , electrophoretic mobility shift ( 46 ) , mass spectrometry ( 3 , 5 , 6 , 7 , 8 , 9 , 10 , 11 , 12 , 13 , 14 , 15 , 16 , 17 , 18 , 19 , 20 , 21 , 22 , 23 , 24 , 25 , 26 , 27 , 28 , 29 , 30 , 31 , 32 , 33 , 34 , 35 , 36 , 37 , 38 , 39 ) , mass spectrometry (in vitro) ( 13 ) , mutation of modification site ( 42 ) , peptide sequencing ( 46 ) , phospho-antibody ( 1 , 2 , 40 , 41 , 42 ) , western blotting ( 1 , 2 , 40 , 41 , 42 )
Disease tissue studied:	anthrax infection ( 17 ) , neuroblastoma ( 16 ) , melanoma skin cancer ( 29 ) , testicular cancer ( 24 )
Relevant cell line - cell type - tissue:	'fat, brown' ( 18 ) , 'fat, brown'-'fat, brown' ( 19 ) , 3T3 (fibroblast) [CDC42 (human), transfection] ( 7 ) , 3T3 (fibroblast) [KRas (human), transfection] ( 7 ) , 3T3 (fibroblast) ( 7 ) , astrocyte ( 2 ) , brain ( 20 ) , breast cell ( 46 ) , C2C12 (myoblast) ( 13 , 19 , 30 , 31 , 32 , 33 , 34 , 39 ) , ES-J1 (stem) ( 23 , 25 ) , F9 (testicular) ( 24 ) , heart ( 9 , 18 , 35 , 36 , 37 ) , HL-1 (myocyte) [Akt1 (mouse), knockdown, stable lentiviral expression of Akt1 shRNA] ( 5 ) , HL-1 (myocyte) [Akt2 (mouse), knockdown, stable lentiviral expression of Akt2 shRNA] ( 5 ) , HL-1 (myocyte) ( 5 ) , kidney ( 18 ) , L929 (fibroblast) ( 42 ) , liver ( 3 , 8 , 11 , 18 , 21 , 22 , 27 , 28 , 38 ) , liver [leptin (mouse), homozygous knockout] ( 11 ) , lung ( 18 ) , macrophage-peritoneum ( 10 ) , MEF (fibroblast) [p53 (mouse), homozygous knockout] ( 14 ) , MEF (fibroblast) [TSC2 (mouse), homozygous knockout] ( 15 ) , MEF (fibroblast) ( 10 , 40 ) , muscle ( 26 ) , myocyte-heart ( 1 , 41 ) , N1E-115 (neuron) ( 16 ) , pancreas ( 18 ) , PC-12 (chromaffin) [TrkA (rat), transfection] ( 12 ) , PC-12 (chromaffin) ( 12 ) , skin [mGluR1 (mouse), transgenic, TG mutant mice] ( 29 ) , spleen ( 17 , 18 ) , stromal ( 6 ) , testis ( 18 )

Upstream Regulation
Regulatory protein:	CDC42 (mouse) ( 7 ) , CYBB (human) ( 1 ) , KRas (mouse) ( 7 ) , MAPKAPK2 (mouse) ( 40 ) , P2X7 (mouse) ( 2 ) , p47phox (human) ( 1 ) , RIPK3 (mouse) ( 10 ) , TSC2 (mouse) ( 14 )
Putative in vivo kinases:	PKACA (mouse) ( 46 ) , PKCA (mouse) ( 46 ) , PKCD (mouse) ( 42 )
Kinases, in vitro:	MAPKAPK2 (human) ( 45 ) , PKACA (mouse) ( 46 ) , PKCA (mouse) ( 46 ) , PKCD (mouse) ( 42 )
Treatments:	arsenite ( 40 ) , FI14 ( 2 ) , geldanamycin ( 1 ) , GS-444217 ( 1 ) , H2O2 ( 40 , 42 ) , heat_shock ( 40 ) , ischemia ( 6 ) , kainic_acid ( 2 ) , midazolam ( 41 ) , OSMI-1 ( 1 ) , PD169316 ( 41 ) , PD98059 ( 43 ) , PDGF ( 12 , 43 ) , phenylephrine ( 1 ) , phorbol_ester ( 42 ) , rottlerin ( 42 ) , SB202190 ( 1 ) , SB202474 ( 40 ) , SB203580 ( 40 , 41 , 43 ) , SP600125 ( 43 ) , thrombin ( 41 )

Downstream Regulation
Effects of modification on HSP27:	activity, induced ( 42 ) , intracellular localization ( 42 )
Effects of modification on biological processes:	apoptosis, inhibited ( 42 )

References
1	Papanicolaou KN, et al. (2023) Inhibiting O-GlcNAcylation impacts p38 and Erk1/2 signaling and perturbs cardiomyocyte hypertrophy. J Biol Chem 299, 102907 36642184 Curated Info
2	Lee DS, Kim JE (2020) P2 ¿¿ 7 Receptor Inhibits Astroglial Autophagy via Regulating FAK- and PHLPP1/2-Mediated AKT-S473 Phosphorylation Following Kainic Acid-Induced Seizures. Int J Mol Sci 32899862 Curated Info
3	Robles MS, Humphrey SJ, Mann M (2017) Phosphorylation Is a Central Mechanism for Circadian Control of Metabolism and Physiology. Cell Metab 25, 118-127 27818261 Curated Info
4	Sacco F, et al. (2016) Glucose-regulated and drug-perturbed phosphoproteome reveals molecular mechanisms controlling insulin secretion. Nat Commun 7, 13250 27841257 Curated Info
5	Reinartz M, Raupach A, Kaisers W, Gödecke A (2014) AKT1 and AKT2 induce distinct phosphorylation patterns in HL-1 cardiac myocytes. J Proteome Res 13, 4232-45 25162660 Curated Info
6	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
7	Gnad F, et al. (2013) Systems-wide Analysis of K-Ras, Cdc42, and PAK4 Signaling by Quantitative Phosphoproteomics. Mol Cell Proteomics 12, 2070-80 23608596 Curated Info
8	Wilson-Grady JT, Haas W, Gygi SP (2013) Quantitative comparison of the fasted and re-fed mouse liver phosphoproteomes using lower pH reductive dimethylation. Methods 61, 277-86 23567750 Curated Info
9	Lundby A, et al. (2013) In vivo phosphoproteomics analysis reveals the cardiac targets of β-adrenergic receptor signaling. Sci Signal 6, rs11 23737553 Curated Info
10	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
11	Grimsrud PA, et al. (2012) A quantitative map of the liver mitochondrial phosphoproteome reveals posttranslational control of ketogenesis. Cell Metab 16, 672-83 23140645 Curated Info
12	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
13	Knight JD, et al. (2012) A novel whole-cell lysate kinase assay identifies substrates of the p38 MAPK in differentiating myoblasts. Skelet Muscle 2, 5 22394512 Curated Info
14	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
15	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
16	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
17	Manes NP, et al. (2011) Discovery of mouse spleen signaling responses to anthrax using label-free quantitative phosphoproteomics via mass spectrometry. Mol Cell Proteomics 10, M110.000927 21189417 Curated Info
18	Huttlin EL, et al. (2010) A tissue-specific atlas of mouse protein phosphorylation and expression. Cell 143, 1174-89 21183079 Curated Info
19	Gnad F, et al. (2010) Evolutionary constraints of phosphorylation in eukaryotes, prokaryotes, and mitochondria. Mol Cell Proteomics 9, 2642-53 20688971 Curated Info
20	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
21	Possemato A (2010) CST Curation Set: 9727; Year: 2010; Biosample/Treatment: tissue, liver/AICAR; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: RXXp[ST] Antibodies Used to Purify Peptides prior to LCMS: Phospho-(Ser/Thr) PKD Substrate Antibody Cat#: 4381, PTMScan(R) Phospho-PKD Substrate Motif (LXRXXpS/pT) Immunoaffinity Beads Cat#: 1986 Curated Info
22	Possemato A (2010) CST Curation Set: 9729; Year: 2010; Biosample/Treatment: tissue, liver/AICAR; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: RXXp[ST] Antibodies Used to Purify Peptides prior to LCMS: Phospho-(Ser/Thr) PKD Substrate Antibody Cat#: 4381, PTMScan(R) Phospho-PKD Substrate Motif (LXRXXpS/pT) Immunoaffinity Beads Cat#: 1986 Curated Info
23	Zhou J (2010) CST Curation Set: 9664; Year: 2010; Biosample/Treatment: cell line, ES J1/untreated; Disease: -; 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
24	Zhou J (2010) CST Curation Set: 9671; Year: 2010; Biosample/Treatment: cell line, F9/untreated; Disease: testicular cancer; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: RXXp[ST] Antibodies Used to Purify Peptides prior to LCMS: Phospho-(Ser/Thr) PKD Substrate Antibody Cat#: 4381, PTMScan(R) Phospho-PKD Substrate Motif (LXRXXpS/pT) Immunoaffinity Beads Cat#: 1986 Curated Info
25	Zhou J (2010) CST Curation Set: 9670; Year: 2010; Biosample/Treatment: cell line, ES J1/untreated; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: RXXp[ST] Antibodies Used to Purify Peptides prior to LCMS: Phospho-(Ser/Thr) PKD Substrate Antibody Cat#: 4381, PTMScan(R) Phospho-PKD Substrate Motif (LXRXXpS/pT) Immunoaffinity Beads Cat#: 1986 Curated Info
26	Guo A (2009) CST Curation Set: 8163; Year: 2009; Biosample/Treatment: tissue, muscle/untreated; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: (F/Y/M)Xp[ST](L/I/M) Curated Info
27	Possemato A (2009) CST Curation Set: 8043; Year: 2011; Biosample/Treatment: tissue, liver/AICAR; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: RXXp[ST] Curated Info
28	Possemato A (2009) CST Curation Set: 8042; Year: 2011; Biosample/Treatment: tissue, liver/AICAR; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: RXXp[ST] Curated Info
29	Zanivan S, et al. (2008) Solid tumor proteome and phosphoproteome analysis by high resolution mass spectrometry. J Proteome Res 7, 5314-26 19367708 Curated Info
30	Possemato A (2008) CST Curation Set: 5259; Year: 2008; Biosample/Treatment: cell line, C2C12/serum starved; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: RXXp[ST] Curated Info
31	Possemato A (2008) CST Curation Set: 5027; Year: 2008; Biosample/Treatment: cell line, C2C12/untreated; Disease: -; 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
32	Possemato A (2008) CST Curation Set: 5028; Year: 2008; Biosample/Treatment: cell line, C2C12/untreated; Disease: -; 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
33	Possemato A (2008) CST Curation Set: 5033; Year: 2008; Biosample/Treatment: cell line, C2C12/untreated; Disease: -; 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
34	Possemato A (2008) CST Curation Set: 5034; Year: 2008; Biosample/Treatment: cell line, C2C12/untreated; Disease: -; 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 (2008) CST Curation Set: 3820; Year: 2008; Biosample/Treatment: tissue, heart/untreated; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: RXXp[ST] Antibodies Used to Purify Peptides prior to LCMS: Phospho-(Ser/Thr) PKD Substrate Antibody Cat#: 4381, PTMScan(R) Phospho-PKD Substrate Motif (LXRXXpS/pT) Immunoaffinity Beads Cat#: 1986 Curated Info
36	Guo A (2007) CST Curation Set: 2922; Year: 2007; Biosample/Treatment: tissue, heart/-; 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
37	Guo A (2007) CST Curation Set: 2923; Year: 2007; Biosample/Treatment: tissue, heart/-; Disease: -; SILAC: -; Specificities of Antibodies Used to Purify Peptides prior to LCMS: p[ST](D/E)X(D/E) Antibodies Used to Purify Peptides prior to LCMS: Phospho(Ser/Thr) CKII Substrate Antibody (polyAb) Cat#: 5808, PTMScan(R) Phospho-CK Substrate Motif (S/TD/EXD/E) Immunoaffinity Beads Cat#: 1994 Curated Info
38	Villén J, Beausoleil SA, Gerber SA, Gygi SP (2007) Large-scale phosphorylation analysis of mouse liver. Proc Natl Acad Sci U S A 104, 1488-93 17242355 Curated Info
39	Puente LG, Voisin S, Lee RE, Megeney LA (2006) Reconstructing the regulatory kinase pathways of myogenesis from phosphopeptide data. Mol Cell Proteomics 5, 2244-51 16971385 Curated Info
40	Vertii A, Hakim C, Kotlyarov A, Gaestel M (2006) Analysis of properties of small heat shock protein Hsp25 in MAPK-activated protein kinase 2 (MK2)-deficient cells: MK2-dependent insolubilization of Hsp25 oligomers correlates with susceptibility to stress. J Biol Chem 281, 26966-75 16840785 Curated Info
41	Tanabe K, et al. (2005) Midazolam suppresses thrombin-induced heat shock protein 27 phosphorylation through inhibition of p38 mitogen-activated protein kinase in cardiac myocytes. J Cell Biochem 96, 56-64 16052527 Curated Info
42	Lee YJ, et al. (2005) HSP25 inhibits protein kinase C delta-mediated cell death through direct interaction. J Biol Chem 280, 18108-19 15731106 Curated Info
43	Takenaka M, et al. (2004) Platelet-derived growth factor-BB phosphorylates heat shock protein 27 in cardiac myocytes. J Cell Biochem 91, 316-24 14743391 Curated Info
44	Rogalla T, et al. (1999) Regulation of Hsp27 oligomerization, chaperone function, and protective activity against oxidative stress/tumor necrosis factor alpha by phosphorylation. J Biol Chem 274, 18947-56 10383393 Curated Info
45	Stokoe D, et al. (1992) Identification of MAPKAP kinase 2 as a major enzyme responsible for the phosphorylation of the small mammalian heat shock proteins. FEBS Lett 313, 307-13 1332886 Curated Info
46	Gaestel M, et al. (1991) Identification of the phosphorylation sites of the murine small heat shock protein hsp25. J Biol Chem 266, 14721-4 1860870 Curated Info