A Reference Guide for Jak/STAT Signaling

The Jak/STAT pathway is one of the primary signaling pathways activated by cytokines and growth factors. It is activated by numerous compounds, and each ligand/receptor complex can activate multiple Jak and STAT proteins. To help sort out the complexity of Jak/STAT activation, a list of receptors and the Jak and STAT family members they activate have been assembled into a reference table.

The Jak/STAT Pathway

The Jak/STAT pathway mediates the effects of a large number of cytokines and growth factors. These proteins generally bind to receptors that lack intrinsic kinase activity and thus utilize intracellular enzymes to initiate downstream signaling. This signaling pathway is relatively straightforward as it includes only two main components: Janus Kinases (Jaks) and Signal Transducers and Activators of Transcription (STATs). Jak proteins are non-receptor tyrosine kinases that are constitutively associated with the cytoplasmic domain of receptors. The Jak family includes Jak1, Jak2, Jak3, and Tyk2 (Tyrosine Kinase 2). STAT proteins are latent transcription factors that reside in the cytoplasm until activated via phosphorylation, at which time they translocate to the nucleus. Seven STAT family members have thus far been identified: STAT1, STAT2, STAT3, STAT4, STAT5a, STAT5b, and STAT6.

The Jak/STAT pathway is activated following binding of a cytokine or growth factor to its respective receptor. Receptor engagement by a ligand typically initiates oligomerization of the receptor chains allowing the associated Jak proteins to activate each other by reciprocal tyrosine phosphorylation. The activated Jaks subsequently phosphorylate specific tyrosine residues in the intracellular domains of the receptor. These phosphorylated residues serve as docking sites for the Src homology 2 (SH2)-domain of STAT proteins. Once bound to the receptor, STAT proteins are phosphorylated by Jaks or other kinases, such as Src. Phosphorylated STATs dissociate from the receptor, dimerize, and then translocate to the nucleus where they complex with other nuclear proteins and bind to specific sequences to regulate the expression of target genes.

To date, over 50 cytokines and growth factors have been shown to utilize the four Jak proteins and seven STAT molecules to regulate cell growth, survival, differentiation, motility, and immune responses.1 Activators of the Jak/STAT pathway include type I/II Interferons, the IL-6, IL-10, IL-12, Common gamma Chain, and Common beta Chain families of cytokines, and numerous growth factors that signal through homodimeric receptors, receptor tyrosine kinases, and G protein-coupled receptors. The different receptor classes that activate the Jak/STAT pathway display specificity for one Jak or Jak combination, though they appear to activate multiple STAT proteins.2 Likewise, it does not appear that activated Jaks exhibit specificity for a particular STAT because different receptors activate a common STAT but distinctly different Jaks.3 It is generally accepted, though, that receptors preferably signal through specific Jak and STAT proteins. However, several factors, such as expression profiles of Jak and STAT proteins, the presence of regulatory molecules, and crosstalk from other signaling pathways, can influence which Jak and STAT proteins are activated by a receptor.

The reference table below is a comprehensive index of ligands and receptors, and the Jak and STAT proteins they have been documented to activate. Explore this table to see which Jak and STAT proteins are activated by your receptor of interest.

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Type I/II Interferons

Ligand Receptor Complex Activated Jak Family Member Citation # Activated STAT Family Member Citation #
IFN-alpha IFN-alpha/beta R1 + IFN-alpha/beta R2 Jak1 4 STAT1 13
    Jak2 5 STAT2 13
    Tyk2 6 STAT3 14
        STAT4 15
        STAT5 16
        STAT6 17
IFN-beta IFN-alpha/beta R1 + IFN-alpha/beta R2 Jak1 7 STAT1 7
    Jak2 8 STAT2 7
    Tyk2 9 STAT3 18
        STAT4 19
        STAT5 20
        STAT6 21
IFN-gamma IFN-gamma R1/CD119 + IFN-gamma R2 Jak1 10 STAT1 13
    Jak2 11 STAT2 22
    Tyk2 12 STAT3 14
        STAT5 12
        STAT6 23

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Receptor Tyrosine Kinases

Ligand Receptor
Complex		 Activated Jak Family Member Citation # Activated STAT Family Member Citation #
PBEF/Vistafin Insulin R/CD220 Jak2 24 STAT3 43
IGF-I IGF-I R Jak1 25 STAT3 44
    Jak2 25 STAT5 45
EGF EGF R/ErbB1 Jak1 26 STAT1 46
    Jak2 27 STAT3 47
      12 STAT5 27
EGF ErbB2/Her2 Jak2 28 STAT1 48
        STAT3 28
        STAT5 49
EGF ErbB4/Her4 Jak2 29 STAT1 50
        STAT5 49
PDGF PDGF R alpha + PDGF R beta Jak1 30 STAT1 30
    Jak2 31 STAT3 31
    Tyk2 31 STAT5 51
        STAT6 51
SCF/c-kit Ligand SCF R/c-kit Jak2 32 STAT1 52
        STAT3 53
        STAT5 54
M-CSF M-CSF R Jak1 33 STAT1 33
    Tyk2 33 STAT3 33
        STAT5 55
FGF FGF R1 Jak1 34 STAT1 56
  FGF R2 Jak2 35 STAT3 56
  FGF R3 Jak3 36 STAT5 57
  FGF R4 Tyk2 34   3
Ephrin-A EphA4 Jak2 37 STAT1 37
        STAT3 37
BDNF TrkB Jak2 38 STAT1 38
        STAT3 38
Angiopoietin-2 Tie-2     STAT3 58
        STAT5 59
VEGF VEGF R1/
Flt-1		 Jak2 39 STAT1 60
  VEGF R2/KDR/Flk-1 Tyk2 40 STAT3 60
  VEGF R3/Flt-4     STAT5 39
        STAT6 40
Gas6 Mer     STAT3 61
        STAT5 62
        STAT6 63
HGF HGF R/c-MET Jak1 41 STAT1 64
    Jak2 42 STAT3 65
        STAT5 64

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Homodimeric Hormone Receptors

Ligand Receptor Complex Activated Jak Family Member Citation # Activated STAT Family Member Citation #
Growth Hormone Growth
Hormone R		 Jak1 66 STAT1 75
    Jak2 67 STAT3 66
        STAT5 66
Thrombopoietin/
Tpo		 Thrombopoietin R/ Tpo R Jak2 68 STAT1 69
    Tyk2 69 STAT3 76
        STAT5 76
Erythropoietin Erythropoietin R Jak2 70 STAT1 77
    Jak3 71 STAT3 78
        STAT5 79
Prolactin Prolactin R Jak1 72 STAT1 80
    Jak2 73 STAT3 81
        STAT5 82

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Common beta Chain Receptor Family

Ligand Receptor
Complex		 Activated Jak Family Member Citation # Activated STAT Family Member Citation #
IL-3 IL-3 R alpha/ CD123 + Jak1 76 STAT1 87
  Common beta Chain Jak2 86 STAT3 87
    Tyk2 87 STAT5 91
        STAT6 79
IL-5 IL-5 R alpha/ CD125 + Jak1 88 STAT3 92
  Common beta Chain Jak2 89 STAT5 88
GM-CSF GM-CSF R alpha + Jak1 88 STAT1 93
  Common beta Chain Jak2 90 STAT3 93
        STAT5 91
        STAT6 94

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Common gamma Chain Receptor Family

Ligand Receptor Complex Activated Jak Family Member Citation # Activated STAT Family Member Citation #
IL-2 IL-2 R alpha + IL-2 R beta + Common gamma Chain/IL-2 R gamma Jak1 95 STAT1 107
    Jak2 96 STAT3 108
    Jak3 97 STAT4 109
        STAT5 110
        STAT6 79
IL-4 IL-4 R alpha + Common gamma Chain/IL-2 R gamma Jak1 95 STAT1 111
  IL-4 R alpha + IL-13 R alpha 1 Jak2 98 STAT3 112
    Jak3 99 STAT4 113
    Tyk2 98 STAT5 114
        STAT6 79
IL-7 IL-7 R alpha/CD127 + Common gamma Chain/IL-2 R gamma Jak1 95 STAT1 100
    Jak3 100 STAT2 115
        STAT3 116
        STAT5 116
        STAT6 115
IL-9 IL-9 R + Common gamma Chain/IL-2 R gamma Jak1 95 STAT1 101
    Jak3 95 STAT3 117
    Tyk2 101 STAT5 118
IL-13* IL-13 R alpha 1 + IL-4 R alpha Jak1 98 STAT1 119
    Jak2 98 STAT3 112
    Jak3 102 STAT5 112
    Tyk2 98 STAT6 98
IL-15 IL-15 R alpha + IL-2 R beta + Jak1 103 STAT1 120
  Common gamma Chain/IL-2 R gamma Jak2 104 STAT2 115
    Jak3 103 STAT3 103
    Tyk2 105 STAT4 120
        STAT5 103
        STAT6 115
IL-21 IL-21 R + Common gamma Chain/IL-2 R gamma Jak1 106 STAT1 106
    Jak3 106 STAT3 106
        STAT4 120
        STAT5 121
TSLP** IL-7 R alpha/CD127 + TSLP R Jak1 74 STAT1 83
    Jak2 74 STAT3 84
        STAT5 85
*IL-13 is often associated with the Common gamma Chain receptor family because it binds to the same receptor complex and possesses similar biologic properties as IL-4.

**TSLP is considered to be a IL-7-related cytokine as it shares several functions and a receptor subunit with IL-7.

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IL-6 Family

Ligand Receptor Complex Activated Jak Family Member Citation # Activated STAT Family Member Citation #
IL-6 IL-6 R alpha + gp130 Jak1 10 STAT1 135
    Jak2 122 STAT3 136
    Tyk2 10    
IL-11 IL-11 R alpha + gp130 Jak1 10 STAT1 137
    Jak2 123 STAT3 138
    Tyk2 124    
CNTF CNTF R alpha + LIF R alpha + gp130 Jak1 10 STAT1 139
    Jak2 125 STAT3 138
    Tyk2 124    
Cardiotrophin-1/ CT-1 CT-1 R alpha + LIF R alpha + gp130 Jak1 126 STAT1 140
    Jak2 126 STAT3 126
    Tyk2 126 STAT5 141
LIF LIF R alpha + gp130 Jak1 10 STAT1 142
    Jak2 123 STAT3 136
    Tyk2 127 STAT5 143
Oncostastin M/ OSM OSM R beta + gp130 Jak1 10 STAT1 144
  LIF R alpha + gp130 Jak2 123 STAT3 138
    Tyk2 124 STAT5 145
        STAT6 146
G-CSF*** G-CSF R/CD114 Jak1 128 STAT1 147
    Jak2 129 STAT3 147
    Tyk2 12 STAT5 12
Leptin/OB*** Leptin R Jak1 130 STAT1 148
    Jak2 131 STAT3 131
        STAT5 149
        STAT6 149
IL-31 IL-31 RA + OSM R beta Jak1 132 STAT1 132
    Jak2 133 STAT3 132
        STAT5 132
CLF/CLC CNTF R alpha + LIF R alpha + gp130 Jak1 134 STAT1 150
    Jak2 134 STAT3 134
    Tyk2 134    
***G-CSF and Leptin/OB are often associated with the IL-6 family of cytokines because the G-CSF and Leptin receptors are structurally similar to gp130.

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IL-10 Family

Ligand Receptor Complex Activated Jak Family Member Citation # Activated STAT Family Member Citation #
IL-10 IL-10 R alpha + IL-10 R beta Jak1 151 STAT1 91
    Jak2 152 STAT3 151
    Tyk2 151 STAT5 159
IL-19 IL-20 R alpha + IL-20 R beta Jak1 153 STAT3 160
    Jak2 153    
IL-20 IL-20 R alpha + IL-20 R beta Jak2 154 STAT3 160
  IL-22 R alpha 1 + IL-20 R beta     STAT5 154
IL-22 IL-22 R alpha 1 + IL-10 R beta Jak1 155 STAT1 155
    Tyk2 155 STAT3 155
        STAT5 155
IL-24 IL-20 R alpha + IL-20 R beta Jak1 156 STAT1 161
  IL-22 R alpha 1 + IL-20 R beta     STAT3 156
IL-26/AK155 IL-20 R alpha + IL-10 R beta     STAT1 162
        STAT3 162
IL-28A/IFN-lambda 2 IL-10 R beta + IL-28 R alpha/IFN-lambda R1 Jak2 157 STAT1 163
        STAT2 164
        STAT3 164
IL-28B/IFN-lambda 3 IL-10 R beta + IL-28 R alpha/IFN-lambda R1     STAT1 163
        STAT2 165
IL-29/IFN-lambda 1 IL-10 R beta + IL-28 R alpha/IFN-lambda R1 Jak1 158 STAT1 163
    Tyk2 158 STAT2 163
        STAT3 163
        STAT4 166
        STAT5 166

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IL-12 Family

Ligand Receptor Complex Activated Jak Family Member Citation # Activated STAT Family Member Citation #
IL-12 IL-12 R beta 1 + IL-12 R beta 2 Jak2 167 STAT1 171
    Tyk2 167 STAT3 172
        STAT4 172
        STAT5 173
        STAT6 174
IL-23 IL-12 R beta 1 +
IL-23 R		 Jak2 168 STAT1 168
    Tyk2 168 STAT3 168
        STAT4 168
        STAT5 168
IL-27 IL-27 R alpha/WSX-1/ TCCR + gp130 Jak1 169 STAT1 169
    Jak2 170 STAT2 170
    Tyk2 170 STAT3 170
        STAT4 175
        STAT5 170
        STAT6 176
IL-35 IL-12 R beta 2 + gp130     STAT1 177
        STAT4 177

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G Protein-Coupled Receptors

Ligand Receptor
Complex		 Activated Jak Family Member Citation # Activated STAT Family Member Citation #
Angiotensin II AGTR-1 Jak1 178 STAT1 179
    Jak2 179 STAT2 179
    Tyk2 179 STAT3 195
        STAT5 196
        STAT6 197
Serotonin (5-HT) 5-HT2A Jak1 180 STAT1 180
    Jak2 181 STAT3 181
Coagulation Factor II/ Thrombin PAR1 Jak1 182 STAT1 198
  PAR3 Jak2 183 STAT3 199
  PAR4 Tyk2 182 STAT5 200
Bradykinin Bradykinin RB2/BDKRB2 Jak2 184 STAT3 185
    Tyk2 185    
Platelet-activating Factor (PAF) PAFR Jak2 186 STAT1 187
    Tyk2 187 STAT2 187
        STAT3 187
        STAT5 201
Catecholamines alpha-1A Adrenergic R/ ADRA1A Jak2 188 STAT1 188
  alpha-1B Adrenergic R/ ADRA1B Tyk2 188 STAT3 202
  alpha-1D Adrenergic R/ ADRA1D        
CXCL12/SDF-1 CXCR4 Jak1 189 STAT1 190
    Jak2 190 STAT2 190
    Jak3 190 STAT3 190
    Tyk2 189 STAT4 189
        STAT5 190
CCL2/JE/MCP-1 CCR2 Jak2 191 STAT1 194
        STAT3 191
        STAT5 203
RANTES/CCL5 CCR5 Jak1 192 STAT1 204
    Jak2 193 STAT3 204
    Jak3 193 STAT5 192
CCL15/MIP-1 delta CCR1 Jak1 194 STAT1 194
        STAT3 194

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