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. 2020 Jun 29;9:F1000 Faculty Rev-653. [Version 1] doi: 10.12688/f1000research.22092.1

Table 4. Substrates of p38 group members – others.

Substrate Kinase Function References
Cell-cycle
regulators		 Cdc25A p38α Increases protein stability Goloudina A et al. , Cell Cycle, 2003 133
Cdc25B p38α Increases protein stability Lemaire M et al., Cell Cycle, 2006 134
Cyclin D1 p38α Causes ubiquitination and degradation
of cyclin D1		 Casanovas O et al., J Biol Chem , 2000 135
Cyclin D3 p38α, p38β
p38γ, p38δ		 Causes ubiquitination and degradation
of cyclin D3		 Casanovas O et al., Oncogene, 2004 136
p57kip2 p38α Enhances interaction with CDKs and
inhibits CDKs		 Joaquin M et al., EMBO J, 2012 137
Cell-death
regulators		 Bax p38α Prevents Bcl-2–Bax heterodimer
formation, enhances apoptosis		 Min H et al., Mol Carcinog, 2012 138
BimEL p38α Enhances apoptosis Cai B et al., J Biol Chem, 2006 139
Caspase-3 p38α Inhibits caspase-3 activity and
apoptosis		 Alvarado-Kristensson M et al., J Exp Med,
2004 140
Caspase-8 p38α Inhibits caspase-8 activity and
apoptosis		 Alvarado-Kristensson M et al., J Exp Med,
2004 140
Caspase-9 p38α Inhibits caspase-9 activity and
apoptosis		 Seifert A et al., Cell Signal, 2009 141
DNA/RNA
binding proteins		 Cdt1 p38α, p38β Increases protein stability Chandrasekaran S et al. , Mol Cell Biol,
2011 142
Drosha p38α Enhances nuclear export and
degradation		 Yang Q et al. , Mol Cell, 2015 143
FBP2 p38α Promotes prothrombin mRNA 3' end
processing		 Danckwardt S et al. , Mol Cell, 2011 144
FBP3 p38α Promotes prothrombin mRNA 3' end
processing		 Danckwardt S et al. , Mol Cell, 2011 144
H2AX p38α, p38β Promotes serum starvation-induced
apoptosis		 Lu C et al. , FEBS Lett, 2008 145
H3 p38α N/D Zhong SP et al. , J Biol Chem, 2000 146
HuR p38α, p38β Enhances cytoplasmic accumulation
and increases mRNA stability		 Lafarga V et al. , Mol Cell Biol, 2009 147
KSRP p38α, p38β Prevents KSRP-mediated ARE-directed
mRNA decay		 Briata P et al. , Mol Cell, 2005 148
Rps27 p38α N/D Knight JD et al., Skelet Muscle, 2012 149
SPF45 p38α Inhibits Fas alternative splicing (exon 6
exclusion)		 Al-Ayoubi AM et al., Mol Cell Biol, 2012 150
Endocytosis
regulators		 EEA1 p38α Promotes recruitment to endocytic
membranes and enhances MOR
endocytosis		 Macé G et al. , EMBO J, 2005 151
Rabenosyn-5 p38α Promotes recruitment to endocytic
membranes and enhances MOR
endocytosis		 Macé G et al. , EMBO J, 2005 151
GDI-2 p38α Enhances GDI:Rab5 complex formation
and modulates endocytosis		 Cavalli V et al. , Mol Cell, 2001 152
MAPK pathway
regulator		 JIP4 p38α Enhances p38 activity Kelkar N et al. , Mol Cell Biol, 2005 153
Tip60 p38α Enhances the pro-senescent function
of Tip60		 Zheng H et al. , Mol Cell, 2013 115
TAB1 p38α Inhibits TAK1 activity Cheung PC et al. , EMBO J, 2003 154
TAB3 p38α Inhibits TAK1 activity Mendoza H et al. , Biochem J, 2008 155
FRS2 p38α Downregulates FGF1-induced signaling Zakrzewska M et al., Int J Mol Sci, 2019 156
Membrane
proteins		 EGFR p38α Induces EGFR internalization Winograd-Katz SE et al., Oncogene, 2006 157
FGFR1 p38α Regulates translocation of exogenous
FGF1 into the cytosol/nucleus		 Sørensen V et al., Mol Cell Biol, 2008 158
Nav1.6 p38α Promotes interaction with NEDD-4 and
protein degradation		 Gasser A et al., J Biol Chem, 2010 159
NHE1 p38α Induces intracellular alkalinization Khaled AR et al. , Mol Cell Biol, 2001 160
PLA2 p38α N/D Börsch-Haubold AG et al. , J Biol Chem, 1998 161
TACE p38α, p38β Increases TACE-mediated ectodomain
shedding and TGF-alpha family ligand
release		 Xu P et al. , Mol Cell, 2010 162
ZAP70 p38α Phosphorylation of ZAP70 increases
stability of T cell receptor		 Giardino Torchia ML et al. , Proc Natl Acad Sci
U S A, 2018 41
Structure
proteins		 Caldesmon p38α N/D Hedges JC et al., Am J Physiol, 1998 163
Hsp27 p38α N/D Knight JD et al., Skelet Muscle, 2012 149
Keratin 8 p38α Regulates cellular keratin filament
reorganization		 Ku NO et al., J Biol Chem, 2002 164
Lamin B1 p38α Enhances lamin B1 accumulation Barascu A et al., EMBO J, 2012 165
Paxillin p38α Required for NGF-induced neurite
extension of PC-12 cells		 Huang C et al., J Cell Biol, 2004 166
Stathmin p38δ N/D Parker CG et al., Biochem Biophys Res
Commun, 1998 167
SAP97 p38γ Modulating the association of this
protein with other cytoskeleton proteins		 Sabio G et al., EMBO J, 2005 168
Tau p38α, p38γ, p38δ Enhances formation of paired helical
filaments
Inhibits amyloid-β toxicity in Alzheimer's
mice		 Reynolds CH et al., J Neurochem,1997 169
Ittner A et al., Science, 2016 170
Tensin1 p38α Regulates the binding specificity of
tensin1 to different proteins		 Hall EH et al. , Mol Cell Proteomics, 2010 171
Others DEPTOR p38γ, p38δ Enhances degradation and mTOR
hyperactivation		 González-Terán B et al. , Nat Commun, 2016 114
GS p38β Required for subsequent
phosphorylation to inhibit enzyme activity		 Kuma Y et al. , Biochem J, 2004 172
LAMP2A p38α Activates chaperone-mediated
autophagy		 Li W et al., Nat Commun, 2017 117
Parkin p38α Decreases its interaction with PINK1
and suppresses mitophagy		 Chen J et al., Cell Death Dis, 2018 173
p47 phox p38α Promotes NADPH oxidase activation
and superoxide production		 Makni-Maalej K et al. , J Immunol, 2012 174
p62 p38γ, p38δ Enhances mTORC1 activity Linares JF et al., Cell Rep, 2015 175
Koh A et al., Cell, 2018 176
Raptor p38β Enhances mTORC1 activity in response
to arsenite stress		 Wu X-N et al. , J Biol Chem, 2011 113
Rpn2 p38α Inhibits proteasome activity Lee SH et al. , J Biol Chem, 2010 177
Siah2 p38α Increases Siah2-mediated degradation
of PHD3		 Khurana A et al. , J Biol Chem, 2006 178

CDK, cyclin-dependent kinase; EGFR, epidermal growth factor receptor; FBP1, far upstream binding protein; FGF1, fibroblast growth factor 1; FGFR1, fibroblast growth factor receptor 1; FRS2, fibroblast growth factor receptor substrate 2; GDI, GDP dissociation inhibitor; KSRP, hnRNPK-homology type splicing regulatory protein; MAPK, mitogen-activated protein kinase; mTORC1, mammalian target of rapamycin complex 1; NADPH, nicotinamide adenine dinucleotide phosphate; NGF, nerve growth factor; NHE1, Na +/H + exchanger isoform 1; PHD3, prolyl hydroxylase 3; PLA2, phospholipase A2; SAP97, synapse-associated protein 97; TAB, transforming growth factor-β-activated protein kinase-1-binding protein; TACE, tumor necrosis factor-alpha-converting enzyme; TAK1, transforming growth factor β-activated kinase 1; TGF, transforming growth factor.