Table 1.

Experimentally confirmed platelet-expressed 14-3-3 binding proteins with identified phosphorylated binding sites

UniProtProtein IDProtein nameBinding siteRepresentative functions in cell
P05556 ITB1 Integrin β1 T788 Forms integrin receptors α2β1 and α5β1 for fibronectin, vitronectin, and collagen binding 
P00519 ABL1 Tyrosine-protein kinase ABL1 T735 Cell growth and survival 
Q9UJU6 DBNL Drebrin-like protein T291, S269 Endocytosis, cytoskeleton reorganization 
Q9Y3C5 RNF11 RING finger protein 11 T135 Protein-protein interactions 
Q6R327 RICTR Rapamycin-insensitive companion of mTOR T1135 Cell growth 
O96013 PAK4 Serine/threonine-protein kinase PAK 4 S99 Cytoskeleton reorganization 
Q99683 M3K5 MAPK kinase kinase 5 S966 Cell differentiation and survival 
Q92974 ARHG2 Rho guanine nucleotide exchange factor 2 S886, T114 Multiple cellular processes related to G-protein–coupled receptors 
P16333 NCK1 Cytoplasmic protein NCK1 S85 Cytoplasmic adaptor protein, transducing signals from receptor tyrosine kinases 
P27986 P85A PI3K regulatory subunit α S83 Cytoplasmic adaptor protein/insulin metabolism 
Q99959 PKP2 Plakophilin-2 S82 Protein binding and bridging 
O00159 MYO1C Unconventional myosin-Ic S736 Intracellular movements 
Q8N122 RPTOR Regulatory-associated protein of mTOR S722, S792 Protein bridging/cell growth, survival, and autophagy 
P40818 UBP8 Ubiquitin carboxyl-terminal hydrolase 8 S718 Hydrolysis of ester, thioester, amide, and peptides/regulates protein turnover 
P19634 SL9A1 Sodium/hydrogen exchanger 1 S703 pH regulation/intracellular signal transduction 
Q5PRF9 SMAG2 Protein Smaug homolog 2 S642 Transcription repression 
P07359 GP1BA GPIbα S606, S575 Mediates platelet adhesion and aggregation by binding to VWF A1 domain and other ligands 
Q9Y4H2 IRS2 Insulin receptor substrate 2 S577, S1148 Mediate various cellular processes related to insulin 
Q9NSK0 KLC4 Kinesin light chain 4 S554, S590 Part of kinesin, a microtubule-associated molecular motor 
Q07866 KLC1 Kinesin light chain 1 S545 Part of kinesin, a microtubule-associated molecular motor 
Q06187 BTK Tyrosine-protein kinase BTK S51, T495 Protein and metal ion binding/Cell activation and signaling 
O14974 MYPT1 Protein phosphatase 1, regulatory subunit 12A S472 Part of protein phosphatase 1C 
O60825 F262 6-phosphofructo-2-kinase/fructose-2,6-bisphosphatase 2 S466, S483 Synthesis and degradation of fructose 2,6-bisphosphate 
Q96TC7 RMD3 Regulator of microtubule dynamics protein 3 S46 Calcium homeostasis 
Q9BZL4 PP12C Protein phosphatase 1 regulatory subunit 12C S452 Myosin phosphatase regulator 
Q86VP3 PACS2 Phosphofurin acidic cluster sorting protein 2 S437 Controls endoplasmic reticulum–mitochondria communication/protein trafficking/apoptosis 
Q14432 PDE3A cGMP-inhibited 3′,5′-cyclic phosphodiesterase A S428 Phosphodiesterase activity/cAMP- and cGMP-mediated signaling 
Q7KZI7 MARK2 Serine/threonine-protein kinase MARK2 S400, T596 Protein phosphorylation/microtubule dynamics regulation 
Q13094 LCP2 Lymphocyte cytosolic protein 2 S376 Protein kinase activity regulation/platelet activation 
P15056 BRAF Serine/threonine-protein kinase B-raf S365, S729 Signal transduction 
O15530 PDPK1 3-phosphoinositide-dependent protein kinase 1 S354 Regulates the phosphorylation and activation of a group of protein kinases/signal transduction 
Q9Y4H4 GPSM3 G-protein-signaling modulator 3 S35 Regulates the activation of G(i) α proteins 
Q8IVT5 KSR1 Kinase suppressor of Ras 1 S309, S404 Acts as a scaffold protein that promotes phosphorylation of Raf family members and activation of MAPKs 
Q9Y6R0 NUMBL Numb-like protein S305, S324 Signal transduction 
P23528 COF1 Cofilin-1 S3, S24 Regulates actin cytoskeleton organization 
Q14247 SRC8 Src substrate cortactin S298 Regulates actin cytoskeleton organization and cell deformation/intracellular protein transport 
P48729 KC1A Casein kinase I isoform α S218, S242 Phosphorylates a large number of proteins and participates in a wide variety of cell signaling pathways 
Q9NS28 RGS18 RGS18 S218 Mediates G-protein–coupled receptor signaling pathway 
P10398 ARAF Serine/threonine-protein kinase A-Raf S214, S582 Protein phosphorylation and metal ion binding/cell signaling 
Q8NHG8 ZNRF2 E3 ubiquitin-protein ligase ZNRF2 S19, S82 Protein ubiquitination and ubiquitin-protein transfer 
Q53ET0 CRTC2 CREB-regulated transcription coactivator 2 S171 Glucose homeostasis/cell signaling 
P42575 CASP2 Caspase-2 S164 Apoptosis execution 
Q96F86 EDC3 Enhancer of mRNA-decapping protein 3 S161 mRNA degradation and decapping 
Q13363 CTBP1 C-terminal-binding protein 1 S158 Histone modification/protein binding and phosphorylation 
Q9GZY8 MFF Mitochondrial fission factor S157 Mediates mitochondrial and peroxisomal fission 
Q12802 AKP13 A-kinase anchor protein 13 S1565 Acts as a scaffold protein/mediates signaling downstream of G-protein–coupled receptors 
Q15418 KS6A1 Ribosomal protein S6 kinase α-1 S154 Protein phosphorylation/cell signal transduction and proliferation 
Q13813 SPTN1 Spectrin α chain, non-erythrocytic 1 S1302 Cytoskeleton movement 
Q9UQQ2 SH2B3 SH2B adapter protein 3 S13, S150 Signal transduction/megakaryocyte development and platelet production 
P49815 TSC2 Tuberin S1211, S1254, S939 Regulates protein kinase activity/mediates signal transduction, endocytosis, and cell proliferation 
Q00536 CDK16 Cyclin-dependent kinase 16 S119 Protein phosphorylation/mediates vesicle-mediated transport processes and exocytosis 
O94921 CDK14 Cyclin-dependent kinase 14 S119 Protein phosphorylation/mediates cell proliferation 
Q07889 SOS1 Son of sevenless homolog 1 S1134, S1161 Regulates phosphorylation of MAPK and Ras to Rac signal transduction 
O43896 KIF1C Kinesin-like protein KIF1C S1092 Molecular motor for the transport of vesicles 
Q96PU5 NED4L E3 ubiquitin-protein ligase NEDD4-like S342, S448 Protein ubiquitination/ion transportation and protein localization 
Q2PPJ7 RGPA2 Ral GTPase-activating protein subunit α-2 T715 Subunit of the heterodimeric RalGAP2 complex 
O43524 FOXO3 Forkhead box protein O3 T32, S253 Cell response and apoptosis/protein and DNA binding 
Q8WYL5 SSH1 Protein phosphatase Slingshot homolog 1 S978, S937, S834 Protein dephosphorylation/actin cytoskeleton organization 
P49796 RGS3 Regulator of G-protein signaling 3 S943 Heterotrimeric G-protein signaling suppression 
P21802 FGFR2 Fibroblast growth factor receptor 2 S782 Cell proliferation, differentiation, migration, and apoptosis 
P35222 CTNB1 Catenin β-1 S552 Regulates Wnt signaling pathway/insulin internalization/protein phosphorylation and ubiquitination/cell adhesion, proliferation, differentiation, and apoptosis 
Q9UBF8 PI4KB Phosphatidylinositol 4-kinase β S294 Protein phosphorylation/signal transduction and endocytosis 
Q9UPA5 BSN Protein bassoon S2851 Metal ion binding/synapse assembly 
Q86TI0 TBCD1 TBC1 domain family member 1 S237, T596 Vesicle and protein trafficking/glucose uptake 
Q15418 KS6A1 Ribosomal protein S6 kinase α-1 S154 Protein phosphorylation/signal transduction/cell proliferation, survival, and differentiation 
Q9H6H4 REEP4 Receptor expression-enhancing protein 4 S152 Microtubule binding 
P08069 IGF1R Insulin-like growth factor 1 receptor S1313 Protein phosphorylation/signal transduction/cell migration and proliferation 
P43405 KSYK Tyrosine-protein kinase SYK S297 Signal transduction downstream of transmembrane receptors/mediates platelet adhesion and activation 
P49757 NUMB Protein numb homolog S276, S295 Cell junction organization and cell migration 
Q9UKF7 PITC1 Cytoplasmic phosphatidylinositol transfer protein 1 S274, S299 Lipid transportation/signal transduction 
O75791 GRAP2 GRB2-related adapter protein 2 S262 Interacts with SLP-76 to regulate NF-AT activation/signal transduction 
UniProtProtein IDProtein nameBinding siteRepresentative functions in cell
P05556 ITB1 Integrin β1 T788 Forms integrin receptors α2β1 and α5β1 for fibronectin, vitronectin, and collagen binding 
P00519 ABL1 Tyrosine-protein kinase ABL1 T735 Cell growth and survival 
Q9UJU6 DBNL Drebrin-like protein T291, S269 Endocytosis, cytoskeleton reorganization 
Q9Y3C5 RNF11 RING finger protein 11 T135 Protein-protein interactions 
Q6R327 RICTR Rapamycin-insensitive companion of mTOR T1135 Cell growth 
O96013 PAK4 Serine/threonine-protein kinase PAK 4 S99 Cytoskeleton reorganization 
Q99683 M3K5 MAPK kinase kinase 5 S966 Cell differentiation and survival 
Q92974 ARHG2 Rho guanine nucleotide exchange factor 2 S886, T114 Multiple cellular processes related to G-protein–coupled receptors 
P16333 NCK1 Cytoplasmic protein NCK1 S85 Cytoplasmic adaptor protein, transducing signals from receptor tyrosine kinases 
P27986 P85A PI3K regulatory subunit α S83 Cytoplasmic adaptor protein/insulin metabolism 
Q99959 PKP2 Plakophilin-2 S82 Protein binding and bridging 
O00159 MYO1C Unconventional myosin-Ic S736 Intracellular movements 
Q8N122 RPTOR Regulatory-associated protein of mTOR S722, S792 Protein bridging/cell growth, survival, and autophagy 
P40818 UBP8 Ubiquitin carboxyl-terminal hydrolase 8 S718 Hydrolysis of ester, thioester, amide, and peptides/regulates protein turnover 
P19634 SL9A1 Sodium/hydrogen exchanger 1 S703 pH regulation/intracellular signal transduction 
Q5PRF9 SMAG2 Protein Smaug homolog 2 S642 Transcription repression 
P07359 GP1BA GPIbα S606, S575 Mediates platelet adhesion and aggregation by binding to VWF A1 domain and other ligands 
Q9Y4H2 IRS2 Insulin receptor substrate 2 S577, S1148 Mediate various cellular processes related to insulin 
Q9NSK0 KLC4 Kinesin light chain 4 S554, S590 Part of kinesin, a microtubule-associated molecular motor 
Q07866 KLC1 Kinesin light chain 1 S545 Part of kinesin, a microtubule-associated molecular motor 
Q06187 BTK Tyrosine-protein kinase BTK S51, T495 Protein and metal ion binding/Cell activation and signaling 
O14974 MYPT1 Protein phosphatase 1, regulatory subunit 12A S472 Part of protein phosphatase 1C 
O60825 F262 6-phosphofructo-2-kinase/fructose-2,6-bisphosphatase 2 S466, S483 Synthesis and degradation of fructose 2,6-bisphosphate 
Q96TC7 RMD3 Regulator of microtubule dynamics protein 3 S46 Calcium homeostasis 
Q9BZL4 PP12C Protein phosphatase 1 regulatory subunit 12C S452 Myosin phosphatase regulator 
Q86VP3 PACS2 Phosphofurin acidic cluster sorting protein 2 S437 Controls endoplasmic reticulum–mitochondria communication/protein trafficking/apoptosis 
Q14432 PDE3A cGMP-inhibited 3′,5′-cyclic phosphodiesterase A S428 Phosphodiesterase activity/cAMP- and cGMP-mediated signaling 
Q7KZI7 MARK2 Serine/threonine-protein kinase MARK2 S400, T596 Protein phosphorylation/microtubule dynamics regulation 
Q13094 LCP2 Lymphocyte cytosolic protein 2 S376 Protein kinase activity regulation/platelet activation 
P15056 BRAF Serine/threonine-protein kinase B-raf S365, S729 Signal transduction 
O15530 PDPK1 3-phosphoinositide-dependent protein kinase 1 S354 Regulates the phosphorylation and activation of a group of protein kinases/signal transduction 
Q9Y4H4 GPSM3 G-protein-signaling modulator 3 S35 Regulates the activation of G(i) α proteins 
Q8IVT5 KSR1 Kinase suppressor of Ras 1 S309, S404 Acts as a scaffold protein that promotes phosphorylation of Raf family members and activation of MAPKs 
Q9Y6R0 NUMBL Numb-like protein S305, S324 Signal transduction 
P23528 COF1 Cofilin-1 S3, S24 Regulates actin cytoskeleton organization 
Q14247 SRC8 Src substrate cortactin S298 Regulates actin cytoskeleton organization and cell deformation/intracellular protein transport 
P48729 KC1A Casein kinase I isoform α S218, S242 Phosphorylates a large number of proteins and participates in a wide variety of cell signaling pathways 
Q9NS28 RGS18 RGS18 S218 Mediates G-protein–coupled receptor signaling pathway 
P10398 ARAF Serine/threonine-protein kinase A-Raf S214, S582 Protein phosphorylation and metal ion binding/cell signaling 
Q8NHG8 ZNRF2 E3 ubiquitin-protein ligase ZNRF2 S19, S82 Protein ubiquitination and ubiquitin-protein transfer 
Q53ET0 CRTC2 CREB-regulated transcription coactivator 2 S171 Glucose homeostasis/cell signaling 
P42575 CASP2 Caspase-2 S164 Apoptosis execution 
Q96F86 EDC3 Enhancer of mRNA-decapping protein 3 S161 mRNA degradation and decapping 
Q13363 CTBP1 C-terminal-binding protein 1 S158 Histone modification/protein binding and phosphorylation 
Q9GZY8 MFF Mitochondrial fission factor S157 Mediates mitochondrial and peroxisomal fission 
Q12802 AKP13 A-kinase anchor protein 13 S1565 Acts as a scaffold protein/mediates signaling downstream of G-protein–coupled receptors 
Q15418 KS6A1 Ribosomal protein S6 kinase α-1 S154 Protein phosphorylation/cell signal transduction and proliferation 
Q13813 SPTN1 Spectrin α chain, non-erythrocytic 1 S1302 Cytoskeleton movement 
Q9UQQ2 SH2B3 SH2B adapter protein 3 S13, S150 Signal transduction/megakaryocyte development and platelet production 
P49815 TSC2 Tuberin S1211, S1254, S939 Regulates protein kinase activity/mediates signal transduction, endocytosis, and cell proliferation 
Q00536 CDK16 Cyclin-dependent kinase 16 S119 Protein phosphorylation/mediates vesicle-mediated transport processes and exocytosis 
O94921 CDK14 Cyclin-dependent kinase 14 S119 Protein phosphorylation/mediates cell proliferation 
Q07889 SOS1 Son of sevenless homolog 1 S1134, S1161 Regulates phosphorylation of MAPK and Ras to Rac signal transduction 
O43896 KIF1C Kinesin-like protein KIF1C S1092 Molecular motor for the transport of vesicles 
Q96PU5 NED4L E3 ubiquitin-protein ligase NEDD4-like S342, S448 Protein ubiquitination/ion transportation and protein localization 
Q2PPJ7 RGPA2 Ral GTPase-activating protein subunit α-2 T715 Subunit of the heterodimeric RalGAP2 complex 
O43524 FOXO3 Forkhead box protein O3 T32, S253 Cell response and apoptosis/protein and DNA binding 
Q8WYL5 SSH1 Protein phosphatase Slingshot homolog 1 S978, S937, S834 Protein dephosphorylation/actin cytoskeleton organization 
P49796 RGS3 Regulator of G-protein signaling 3 S943 Heterotrimeric G-protein signaling suppression 
P21802 FGFR2 Fibroblast growth factor receptor 2 S782 Cell proliferation, differentiation, migration, and apoptosis 
P35222 CTNB1 Catenin β-1 S552 Regulates Wnt signaling pathway/insulin internalization/protein phosphorylation and ubiquitination/cell adhesion, proliferation, differentiation, and apoptosis 
Q9UBF8 PI4KB Phosphatidylinositol 4-kinase β S294 Protein phosphorylation/signal transduction and endocytosis 
Q9UPA5 BSN Protein bassoon S2851 Metal ion binding/synapse assembly 
Q86TI0 TBCD1 TBC1 domain family member 1 S237, T596 Vesicle and protein trafficking/glucose uptake 
Q15418 KS6A1 Ribosomal protein S6 kinase α-1 S154 Protein phosphorylation/signal transduction/cell proliferation, survival, and differentiation 
Q9H6H4 REEP4 Receptor expression-enhancing protein 4 S152 Microtubule binding 
P08069 IGF1R Insulin-like growth factor 1 receptor S1313 Protein phosphorylation/signal transduction/cell migration and proliferation 
P43405 KSYK Tyrosine-protein kinase SYK S297 Signal transduction downstream of transmembrane receptors/mediates platelet adhesion and activation 
P49757 NUMB Protein numb homolog S276, S295 Cell junction organization and cell migration 
Q9UKF7 PITC1 Cytoplasmic phosphatidylinositol transfer protein 1 S274, S299 Lipid transportation/signal transduction 
O75791 GRAP2 GRB2-related adapter protein 2 S262 Interacts with SLP-76 to regulate NF-AT activation/signal transduction 

BTK, Bruton tyrosine kinase; mRNA, messenger RNA; mTOR, mammalian target of rapamycin.

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