TABLE 2

Phosphorylation sites in PKGI substrates

Phosphorylated residues are underlined.

Substrate	Phosphorylation Site	Substrate Sequence	GenBank Accession Number	Species	Reference(s)
Consensus sequence	Ser or Thr	XRRXSXXX
ATP-sensitive K⁺ channel	N.D.	N.D.			Han et al., 2001; Chai and Lin, 2008
Bad (Bcl-2 associated death promoter)	Ser155	LRRMSDEF	AAC15100	Rattus norvegicus	Johlfs and Fiscus, 2010
Battenin (CLN3)	N.D.	N.D.			Michalewski et al., 1998
β3-adrenergic receptor	N.D.	N.D.			Angelone et al., 2008
Ca²⁺ channel (voltage-dependent; Ca_v1.2 L-type) α1C subunit	Ser1928	GRRASFHL			Yang et al., 2007
Ca²⁺ channel (voltage-dependent; Ca_v1.2 L-type) β2 subunit	Ser496	SRGLSRQE			Yang et al., 2007
Ca²⁺ channel, L-type, α1c subunit	Ser533	KSKFSRYW			Jiang et al., 2000
Ca²⁺ channel, L-type, α1c subunit	Ser1371	VKLLSRGE			Jiang et al., 2000
Ca²⁺-activated K⁺ channel (α-subunit, cslo-α)	Ser1072	SKKSSSVH			Fukao et al., 1999
Ca²⁺-sensitive K⁺ channel (Hslo)	N.D.	N.D.			Alioua et al., 1998
Calponin homology-associated smooth muscle protein (CHASM)	Ser301	ERRVSAPS			Borman, et al., 2004
cAMP-dependent protein kinase regulatory subunit type I (PKA RI)	Ser100	RGAISAEV			Geahlen and Krebs, 1980; Hashimoto et al., 1981
cGMP-dependent protein kinase anchoring protein 42 (GKAP42)	Ser106	AQKESREE			Yuasa et al., 2000
cGMP-dependent protein kinase type I-α (PKGIα)	Ser1	SEL			Aitken et al., 1984
cGMP-dependent protein kinase type I-α (PKGIα)	Ser50	LPVPSTHI			Aitken et al., 1984
	Thr58	GPRTTRAQ			Glass and Smith, 1983; Aitken et al., 1984
	Ser64	AQGISAEP			Aitken et al., 1984
	Ser72	QTYRSFHD			Aitken et al., 1984
	Thr84	FRKFTKSE			Aitken et al., 1984
cGMP-dependent protein kinase type I-β (PKGIβ)	Ser63	AQKQSAST	NP_035290	Mus musculus	Francis et al., 1996
cGMP-dependent protein kinase type I-β (PKGIβ)	Ser79	RQAISAEP			Smith et al., 1996
Cysteine-rich protein 2 (CRP2) [also known as cysteine-rich LIM-only protein 4 (CRP4)]^{^a}	Ser104	ERKTSGPP			Huber et al., 2000; Zhang et al., 2007; Schmidtko et al., 2008
Cystic fibrosis transmembrane conductance regulator (CFTR)	Ser660	ERRNSILT			Picciotto et al., 1992
Cystic fibrosis transmembrane conductance regulator (CFTR)	Ser700	KRKNSILN			Picciotto et al., 1992
	Ser737	ERRLSLVP			Picciotto et al., 1992
	Ser768	RRRQSVL			Picciotto et al., 1992
	Ser795	TRKVSLA			Picciotto et al., 1992
	Ser813	SRRLSQET			Picciotto et al., 1992
Formin homology domain-containing protein (FHOD1)	Ser1131	ERKRSRGN	NP_037373	Homo sapiens	Wang et al., 2004
GABA-A receptor	Ser409	RKPLSSRE	NP_000803	H. sapiens	Leidenhemier, 1996; McDonald and Moss, 1997
G-septin	Ser91	SRKASSWN			Xue et al., 2000
G-substrate^{^b}	Thr68	RRKDTPAL		Oryctolagus cuniculus	Aitken et al., 1981; Aswad and Greengard, 1981; Endo et al., 1999
	Thr72	RRKDTPAL		R. norvegicus	Endo et al., 2003
	Thr119	RRKDTPAL		O. cuniculus	Aitken et al., 1981; Aswad and Greengard, 1981; Endo et al., 1999
	Thr123	RRKDTPAL		R. norvegicus	Endo et al., 2003
Guanylyl cyclase, soluble	Ser64	QRKTSRNR			Murthy, 2001; Zhou et al., 2008
27-kDa Heat shock protein (HSP 27)	Thr143	TRKYTLPP	AAAA62175	H. sapiens	Butt et al., 2001
Histone 2B	Ser32	RKRSRKE			Glass and Krebs, 1979; Glass and Krebs, 1982
Histone 2B	Ser36	RKESYSV			Glass and Krebs, 1979
Inositol triphosphate receptor (IP3R)	Ser1589	ARRDSVLA			Komalavilas and Lincoln, 1994; Haug et al., 1999; Murthy and Zhou, 2003; Wagner et al., 2003
	Ser1755	GRRESLTS			Komalavilas and Lincoln, 1994; Haug et al., 1999; Murthy and Zhou, 2003; Wagner et al., 2003
Inositol triphosphate receptor-associated PKG substrate isoform A (IRAGa)#	Ser683	ARSMSLSL			Schlossmann et al., 2000; Geiselhöringer et al., 2004
	Ser696	RRRVSVAV			Schlossmann et al., 2000; Ammendola et al., 2001
IRAGb	Ser644	RRRVSVAV	AAF61203	Bos taurus	Casteel et al., 2008
Large conductance calcium activated potassium channel (BK_Ca)^{^b}	N.D.	N.D.			Sausbier et al., 2000
LIM and SH3 domain protein (LASP-1)	Ser61	YPKQSFTM	AAK28338	R. norvegicus	Keicher et al., 2004
	Thr156	YRRPTEQQ	AAK28338	R. norvegicus	Keicher et al., 2004
	Ser146	ERRDSQDG	NP_006139	H. sapiens	Butt et al., 2003
MEKK1	N.D.	N.D.			Soh et al., 2001
Myosin phosphatase small regulatory subunit (M20)	N.D.	N.D.			Nakamura et al., 1999
Myosin phosphatase large regulatory subunit (MYPT1)^{^b}	Ser695	QSRRSTQG			Nakamura et al., 1999; Wooldridge et al., 2004
Na⁺/K⁺ ATPase α-subunit	N.D.	N.D.			Fotis et al., 1999
NC-1.1 receptor	N.D.	N.D.			Holmgreen et al., 1997
p21-activated kinase 1 (Pak)	Ser21	MRNTSTMI			Fryer et al., 2006
Phosphodiesterase 5^{^b}	Ser92	TRKISASE			Thomas et al., 1990
Phospholamban^{^b}	Ser16	IRRASTIE	NP_002658	H. sapiens	Raeymaekers et al., 1988; Cornwell et al., 1991
Phospholipase A₂ (PLA₂)	N.D.	N.D.			Chalimoniuk et al., 2009
Phospholipase C β₂ (PLCβ₂)	N.D.	N.D.			Xia et al., 2001
Phospholipase C β₃ (PLCβ₃)	Ser26	LRRGSKEI			Xia et al., 2001
	Ser1105	KRHNSISE			Xia et al., 2001
pH-sensitive K⁺ channel	N.D.	N.D.			Kang et al., 2007
Protein phosphatase inhibitor 1 (PPI-1)	Thr35	RRPTATL			Tokui et al., 1996
Rap1 GTP-ase activating protein 2-α (Rap1GAP2α)^{^a}	Ser7	GRKRSVSF	CAF31653	H. sapiens	Schultess et al., 2005
Rap1GAP2β^{^a}	Ser7	GRKRSVSF	CAF31652	H. sapiens	Schultess et al., 2005
Regulator of G protein signaling 2 (RGS2)^{^b}	Ser46	KTRLSYFL			Tang et al., 2003; Sun et al., 2005
	Ser64	GKKSKQQ			Tang et al., 2003; Sun et al., 2005
RGS4	Ser52	QRVSQEE	BAA20400	M. musculus	Huang et al., 2007
RhoA^{^b}	Ser188	GKKKSGCL	AAM21117	H. sapiens	Sawada et al., 2001; Ellerbroek et al., 2003; Murthy and Zhou, 2003
Ryanodine receptor	N.D.	N.D.			Takasago et al., 1991
Septin-3^{^a}	Ser91	SRKASSWN	NP_062248	R. norvegicus	Xue et al., 2004
Serotonin transporter (SERT)^{^a}	Thr276	KGVKTSGK	CAA71909	R. norvegicus	Ramamoorthy et al., 2007; Zhang et al., 2007
Smooth myosin phosphatase light chain phosphatase (SMPP-1M)	N.D.	N.D.			Wu et al., 1996; Lee et al., 1997
Smoothelin-like protein 1 (SMTNL1)	Ser301	RRVSARS	NP_001099035	M. musculus	Wooldridge et al., 2008
Splicing factor 1	Ser20	RKRSRWNQ			Wang et al., 1999
Steroidogenic acute regulatory protein (StAR)	N.D.	N.D.			Andric et al., 2007
Telokin^{^b}	Ser13	GRKSSTGS	AAN63946	M. musculus	Walker et al., 2001
Thromboxane A₂ receptor 1α (Tpα)^{^a}	Ser331	PRSLSLQP			Yamamoto et al., 2001; Reid and Kinsella, 2003
Titin N2BA and N2B	Ser469	GAKTSLQE			Kruger et al., 2009
Transient receptor potential	Thr11	LRRMTVMR	NP_003296	H. sapiens	Kwan et al., 2004
Ca²⁺ channel isoform 3 (TRPC3)&	Ser263	YRKLSMQC	NP_003296	H. sapiens	Kwan et al., 2004
Transient receptor potential	Thr70	HRRQTVLR	NP_004612	H. sapiens	Koitabashi et al., 2010
Ca²⁺ channel isoform 6 (TRPC6)	Ser322	YKKLSMQC	NP_004612	H. sapiens	Koitabashi et al., 2010
Troponin I	N.D.	N.D.			Layland et al., 2002
Tyrosine hydroxylase	N.D.	N.D.			Rodríguez-Pascual et al., 1999
Vasodilator-stimulated phosphoprotein (VASP)	Ser157	ERRVSNAG			Smolenski et al., 1998
Vasodilator-stimulated phosphoprotein (VASP)	Ser239	LRKVSKQE			Butt et al., 1994
	Thr278	RRKATQVG			Cook and Haynes, 2007
Vimentin^{^b}	N.D.	N.D.			Wyatt et al., 1991; Pryzwansky et al., 1995

N.D., not determined or not identified.
↵a PKGI substrates identified in heterologous systems (Hofmann et al., 2009).
↵b Established PKGI substrates (Hofmann et al., 2009).