GAL<sub>1</sub> receptor | Galanin receptors | IUPHAR/BPS Guide to PHARMACOLOGY

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GAL1 receptor

Target not currently curated in GtoImmuPdb

Target id: 243

Nomenclature: GAL1 receptor

Family: Galanin receptors

Annotation status:  image of a green circle Annotated and expert reviewed. Please contact us if you can help with updates.  » Email us

Gene and Protein Information
class A G protein-coupled receptor
Species TM AA Chromosomal Location Gene Symbol Gene Name Reference
Human 7 349 18q23 GALR1 galanin receptor 1 67
Mouse 7 348 18 E3-E4 Galr1 galanin receptor 1 38,98
Rat 7 346 18q12.3 Galr1 galanin receptor 1 15
Previous and Unofficial Names
GALNR | GALNR1
Database Links
Specialist databases
GPCRDB galr1_human (Hs), galr1_mouse (Mm), galr1_rat (Rn)
Other databases
ChEMBL Target
Ensembl Gene
Entrez Gene
Human Protein Atlas
KEGG Gene
OMIM
RefSeq Nucleotide
RefSeq Protein
UniProtKB
Wikipedia
Natural/Endogenous Ligands
galanin {Sp: Human} , galanin {Sp: Mouse, Rat}
galanin-like peptide {Sp: Human} , galanin-like peptide {Sp: Mouse} , galanin-like peptide {Sp: Rat}
Comments: Galanin is more potent than galanin-like peptide
Potency order of endogenous ligands (Human)
galanin (GAL, P22466) > galanin-like peptide (GALP, Q9UBC7)  [68]

Download all structure-activity data for this target as a CSV file

Agonists
Key to terms and symbols View all chemical structures Click column headers to sort
Ligand Sp. Action Value Parameter Reference
[125I][Tyr26]galanin (pig) Rn Agonist 9.5 – 10.7 pKd 15,20,22,71,85-86
pKd 9.5 – 10.7 (Kd 3x10-10 – 1.9x10-11 M) Two populations of binding sites have been identified in transfected cells with Kds of 0.3 and 0.019nM. [15,20,22,71,85-86]
[125I][Tyr26]galanin (rat/mouse) Mm Agonist 9.9 pKd 98
pKd 9.9 (Kd 1.34x10-10 M) [98]
[125I][Tyr26]galanin (pig) Hs Agonist 9.8 pKd 11
pKd 9.8 (Kd 1.5x10-10 M) [11]
[19Lys,26Leu]-galparan Rn Agonist 9.1 pKd 46
pKd 9.1 (Kd 7.1x10-10 M) [46]
galanin {Sp: Pig} Rn Full agonist 9.1 pKd 46
pKd 9.1 (Kd 7.4x10-10 M) [46]
galanin {Sp: Pig} Hs Full agonist 9.1 pKd 29
pKd 9.1 (Kd 8x10-10 M) [29]
[125I][Tyr26]galanin (human) Hs Agonist 7.8 – 10.3 pKd 24
pKd 10.3 (Kd 4.89x10-11 M) High affinity receptors [24]
pKd 7.8 (Kd 1.47x10-8 M) Low affinity receptors [24]
galparan Rn Agonist 8.2 pKd 46
pKd 8.2 (Kd 6.4x10-9 M) [46]
galanin(1-13)amide Rn Agonist 6.9 pKd 46
pKd 6.9 (Kd 1.25x10-7 M) [46]
[2Ala]-galparan Rn Agonist 5.8 pKd 46
pKd 5.8 (Kd 1.585x10-6 M) [46]
galanin {Sp: Pig} Hs Full agonist 9.6 – 10.6 pKi 11,24,29
pKi 9.6 – 10.6 (Ki 2.3x10-10 – 2.5x10-11 M) [11,24,29]
galanin {Sp: Mouse, Rat} Mm Full agonist 9.3 – 10.3 pKi 70,98
pKi 9.3 – 10.3 (Ki 4.6x10-10 – 4.7x10-11 M) [70,98]
galanin {Sp: Human} Hs Full agonist 9.1 – 10.5 pKi 11,24,29,82
pKi 9.1 – 10.5 (Ki 8x10-10 – 3.1x10-11 M) [11,24,29,82]
M617 Hs Agonist 9.6 pKi 55
pKi 9.6 (Ki 2.3x10-10 M) [55]
M242 Hs Agonist 9.6 pKi 82
pKi 9.6 (Ki 2.5x10-10 M) [82]
galanin {Sp: Mouse, Rat} Rn Full agonist 9.0 – 10.1 pKi 15,86,94,96-97
pKi 9.0 – 10.1 (Ki 1x10-9 – 8x10-11 M) [15,86,94,96-97]
galanin {Sp: Pig} Rn Full agonist 9.1 – 10.0 pKi 15,71,85-86
pKi 9.1 – 10.0 (Ki 8.2x10-10 – 1x10-10 M) [15,71,85-86]
galanin {Sp: Human} Rn Full agonist 9.2 – 9.9 pKi 15,86
pKi 9.2 – 9.9 (Ki 6x10-10 – 1.4x10-10 M) [15,86]
galanin {Sp: Mouse, Rat} Hs Full agonist 8.8 – 10.1 pKi 11,29,53,55,78,81,88
pKi 8.8 – 10.1 (Ki 1.75x10-9 – 8x10-11 M) [11,29,53,55,78,81,88]
Gal-(K)4 Hs Agonist 9.4 pKi 106
pKi 9.4 (Ki 4x10-10 M) [106]
Gal-B2-MPEG4 Hs Agonist 9.3 pKi 106
pKi 9.3 (Ki 5x10-10 M) [106]
Gal-B2-C8 Hs Agonist 9.1 pKi 106
pKi 9.1 (Ki 7x10-10 M) [106]
Gal-B2-dPEG24 Hs Agonist 8.9 pKi 105
pKi 8.9 (Ki 1.3x10-9 M) [105]
Gal-B2-C10 Hs Agonist 8.9 pKi 106
pKi 8.9 (Ki 1.3x10-9 M) [106]
Gal-B2-C12 Hs Agonist 8.9 pKi 106
pKi 8.9 (Ki 1.4x10-9 M) [106]
galanin(1-16) (rat/mouse/pig) Mm Agonist 8.7 – 9.0 pKi 70,98
pKi 8.7 – 9.0 (Ki 2.2x10-9 – 1.02x10-9 M) [70,98]
galanin(1-16) (rat/mouse/pig) Hs Agonist 8.3 – 9.3 pKi 11,14,24,29
pKi 8.3 – 9.3 (Ki 5x10-9 – 5x10-10 M) [11,14,24,29]
Gal-B2-C14 Hs Agonist 8.6 pKi 106
pKi 8.6 (Ki 2.6x10-9 M) [106]
galanin(1-16) (rat/mouse/pig) Rn Agonist 8.1 – 9.0 pKi 15,71,85-86,94,96-97
pKi 8.1 – 9.0 (Ki 9x10-9 – 9.5x10-10 M) [15,71,85-86,94,96-97]
Gal-B2 Hs Agonist 8.5 pKi 14
pKi 8.5 (Ki 3.5x10-9 M) [14]
Gal-B2-C18 Hs Agonist 8.4 pKi 106
pKi 8.4 (Ki 4x10-9 M) [106]
galanin(1-15) Rn Agonist 7.9 – 8.5 pKi 71,85
pKi 7.9 – 8.5 (Ki 1.15x10-8 – 3x10-9 M) [71,85]
galanin(7-29) (pig) Hs Agonist 8.2 pKi 11
pKi 8.2 (Ki 6.76x10-9 M) [11]
galanin(2-29) (pig) Rn Full agonist 7.1 – 8.1 pKi 15,71
pKi 7.1 – 8.1 (Ki 8.25x10-8 – 7.14x10-9 M) [15,71]
galanin(7-29) (pig) Rn Agonist 7.6 pKi 86
pKi 7.6 (Ki 2.455x10-8 M) [86]
J20 (galanin analogue) Hs Agonist 7.6 pKi 79
pKi 7.6 (Ki 2.5x10-8 M) [79]
galanin(2-29) (pig) Hs Full agonist 7.6 pKi 11
pKi 7.6 (Ki 2.63x10-8 M) [11]
galanin(2-29) (rat/mouse) Mm Full agonist 7.1 – 8.0 pKi 70,98
pKi 7.1 – 8.0 (Ki 8.46x10-8 – 1.08x10-8 M) [70,98]
galanin(2-30) (human) Hs Agonist 7.3 pKi 17
pKi 7.3 (Ki 5.2x10-8 M) [17]
M1151 Hs Agonist 7.0 pKi 81
pKi 7.0 (Ki 9.86x10-8 M) [81]
J18 (galanin analogue) Hs Agonist 6.9 pKi 79
pKi 6.9 (Ki 1.38x10-7 M) [79]
galanin(2-29) (rat/mouse) Rn Full agonist 6.0 – 7.1 pKi 94,96-97
pKi 6.0 – 7.1 (Ki 1.1x10-6 – 8.5x10-8 M) [94,96-97]
[N-Me,des-Sar]Gal-B2 Hs Agonist 6.4 pKi 77
pKi 6.4 (Ki 3.645x10-7 M) [77]
Gal-B5 Hs Agonist 6.4 pKi 14
pKi 6.4 (Ki 3.87x10-7 M) [14]
M1145 Hs Agonist 6.2 pKi 78
pKi 6.2 (Ki 5.87x10-7 M) [78]
galanin(D-Trp2) (pig) Hs Agonist 6.2 pKi 11
pKi 6.2 (Ki 6.31x10-7 M) [11]
galanin(D-Trp2) (pig) Rn Agonist 6.0 – 6.4 pKi 85-86
pKi 6.0 – 6.4 (Ki 1x10-6 – 4.074x10-7 M) [85-86]
galanin(3-29) (pig) Rn Agonist <6.0 pKi 15,71,85-86
pKi <6.0 (Ki >1x10-6 M) [15,71,85-86]
galanin(3-29) (rat/mouse) Rn Agonist <6.0 pKi 94,97
pKi <6.0 (Ki >1x10-6 M) [94,97]
galanin(10-29) (pig) Rn Agonist <6.0 pKi 71
pKi <6.0 (Ki >1x10-6 M) [71]
galanin(3-29) (rat/mouse) Mm Agonist <5.8 pKi 70
pKi <5.8 (Ki >1.65x10-6 M) [70]
galanin(10-29) (rat/mouse) Mm Agonist <5.8 pKi 70
pKi <5.8 (Ki >1.67x10-6 M) [70]
M1153 Hs Agonist 5.7 pKi 81
pKi 5.7 (Ki 1.89x10-6 M) [81]
M1152 Hs Agonist 5.6 pKi 81
pKi 5.6 (Ki 2.37x10-6 M) [81]
galanin(3-29) (pig) Hs Agonist <5.0 – 6.0 pKi 11,29
pKi <5.0 – 6.0 (Ki >1x10-5 – 1x10-6 M) [11,29]
galanin(2-11) Hs Agonist <5.3 pKi 53
pKi <5.3 (Ki >5x10-6 M) [53]
galnon Hs Agonist 4.9 pKi 5,87
pKi 4.9 (Ki 1.17x10-5 M) [5,87]
M1160 Hs Agonist 4.8 pKi 80
pKi 4.8 (Ki 1.55x10-5 M) [80]
galmic Hs Agonist 4.5 pKi 5,87
pKi 4.5 (Ki 3.42x10-5 M) [5,87]
galanin {Sp: Mouse, Rat} Rn Full agonist 9.8 pEC50 68
pEC50 9.8 (EC50 1.6x10-10 M) [68]
galanin-like peptide {Sp: Pig} Rn Agonist 7.5 pEC50 68
pEC50 7.5 (EC50 3x10-8 M) [68]
galanin {Sp: Human} Rn Full agonist 10.1 – 10.2 pIC50 33,91
pIC50 10.1 – 10.2 (IC50 7x10-11 – 6x10-11 M) [33,91]
galanin {Sp: Pig} Rn Full agonist 10.1 – 10.2 pIC50 33,91
pIC50 10.1 – 10.2 (IC50 7x10-11 – 6x10-11 M) [33,91]
galanin {Sp: Mouse, Rat} Hs Full agonist 9.9 pIC50 22
pIC50 9.9 (IC50 1.3x10-10 M) [22]
galanin {Sp: Pig} Hs Full agonist 9.8 – 9.9 pIC50 22,91
pIC50 9.8 – 9.9 (IC50 1.6x10-10 – 1.4x10-10 M) [22,91]
galanin(1-19) (human) Rn Agonist 9.8 pIC50 91
pIC50 9.8 (IC50 1.5x10-10 M) [91]
galanin {Sp: Human} Hs Full agonist 9.6 – 9.9 pIC50 22,49,91
pIC50 9.6 – 9.9 (IC50 2.7x10-10 – 1.3x10-10 M) [22,49,91]
galanin(1-16) (rat/mouse/pig) Rn Agonist 9.6 pIC50 33,91
pIC50 9.6 (IC50 2.7x10-10 M) [33,91]
galanin(1-19) (human) Hs Agonist 9.5 pIC50 91
pIC50 9.5 (IC50 3.4x10-10 M) [91]
galanin {Sp: Mouse, Rat} Rn Full agonist 8.9 – 10.0 pIC50 13,68
pIC50 8.9 – 10.0 (IC50 1.4x10-9 – 9.7x10-11 M) [13,68]
[Sar1, D-Ala12]galanin(1-16) Hs Agonist 9.4 pIC50 47
pIC50 9.4 (IC50 4x10-10 M) [47]
galanin(1-16) (rat/mouse/pig) Hs Agonist 9.0 – 9.2 pIC50 22,91
pIC50 9.0 – 9.2 (IC50 1.01x10-9 – 6x10-10 M) [22,91]
galanin(1-11) Hs Agonist 9.0 pIC50 49
pIC50 9.0 (IC50 1.1x10-9 M) [49]
galanin(1-15) Hs Agonist 8.8 pIC50 22
pIC50 8.8 (IC50 1.68x10-9 M) [22]
galanin-like peptide {Sp: Pig} Rn Agonist 8.4 pIC50 68
pIC50 8.4 (IC50 4.3x10-9 M) [68]
galanin(2-29) (rat/mouse) Rn Full agonist 8.1 pIC50 33,91
pIC50 8.1 (IC50 7.2x10-9 M) [33,91]
galanin(2-29) (rat/mouse) Hs Full agonist 8.0 pIC50 91
pIC50 8.0 (IC50 1x10-8 M) [91]
galanin(2-29) (pig) Hs Full agonist 7.5 pIC50 22
pIC50 7.5 (IC50 3.04x10-8 M) [22]
GALP(3-32) (human) Hs Agonist 7.5 pIC50 44
pIC50 7.5 (IC50 3.3x10-8 M) [44]
galanin-like peptide {Sp: Rat} Rn Agonist 7.3 pIC50 13
pIC50 7.3 (IC50 4.5x10-8 M) [13]
galanin-like peptide {Sp: Human} Hs Agonist 7.1 pIC50 44
pIC50 7.1 (IC50 7.7x10-8 M) [44]
GALP(1-32) (human) Hs Agonist 6.9 pIC50 44
pIC50 6.9 (IC50 1.29x10-7 M) [44]
galanin(2-11) Hs Agonist 6.1 pIC50 49
pIC50 6.1 (IC50 8.79x10-7 M) [49]
galanin(3-29) (pig) Hs Agonist <6.0 pIC50 22
pIC50 <6.0 (IC50 >1x10-6 M) [22]
galanin(3-29) (rat/mouse) Hs Agonist <6.0 pIC50 91
pIC50 <6.0 (IC50 >1x10-6 M) [91]
galanin(3-29) (rat/mouse) Rn Agonist <6.0 pIC50 33,91
pIC50 <6.0 (IC50 >1x10-6 M) [33,91]
alarin {Sp: Rat} Rn Agonist <6.0 pIC50 13
pIC50 <6.0 (IC50 >1x10-6 M) [13]
View species-specific agonist tables
Agonist Comments
Rat and mouse galanin sequences are identical [54]. Binding affinity is lost with removal of amino acids 1 and 2 of galanin [11].
Galanin(D-Trp2) and galanin(2-29) both have higher binding affinity for GAL2 than GAL1 receptors [11].
Many different galanin analogues were trialled in [14] and [77]. The most active compounds are reported in the table above. The Ki values of other analogues are listed in these publications.

[94] and [24] have data for two types of radioligand binding assays; membrane binding and whole-cell binding assays.

NOTE that compounds with -ve log10 affinities <6 have no appreciable agonist function.
Antagonists
Key to terms and symbols View all chemical structures Click column headers to sort
Ligand Sp. Action Value Parameter Reference
M15 Hs Antagonist 9.4 – 10.6 pKi 11,24,53
pKi 9.4 – 10.6 (Ki 4x10-10 – 2.7x10-11 M) [11,24,53]
M40 Mm Antagonist 9.4 – 10.4 pKi 70,98
pKi 9.4 – 10.4 (Ki 4.3x10-10 – 4x10-11 M) [70,98]
M32 Hs Antagonist 9.6 – 9.7 pKi 11,82
pKi 9.6 – 9.7 (Ki 2.6x10-10 – 1.8x10-10 M) [11,82]
C7 Hs Antagonist 9.2 – 10.1 pKi 11,24
pKi 9.2 – 10.1 (Ki 6.9x10-10 – 8.2x10-11 M) [11,24]
M35 Hs Antagonist 8.3 – 10.9 pKi 11,24,53
pKi 8.3 – 10.9 (Ki 4.8x10-9 – 1.2x10-11 M) [11,24,53]
M35 Rn Antagonist 9.3 – 9.5 pKi 71,85-86
pKi 9.3 – 9.5 (Ki 4.6x10-10 – 3x10-10 M) [71,85-86]
M40 Hs Antagonist 8.6 – 10.0 pKi 11,24,53
pKi 8.6 – 10.0 (Ki 2.4x10-9 – 1.1x10-10 M) [11,24,53]
M32 Rn Antagonist 9.2 – 9.2 pKi 85-86
pKi 9.2 – 9.2 (Ki 6.8x10-10 – 6x10-10 M) [85-86]
C7 Mm Antagonist 8.7 – 9.0 pKi 70,98
pKi 8.7 – 9.0 (Ki 2x10-9 – 9.6x10-10 M) [70,98]
C7 Rn Antagonist 7.8 – 9.6 pKi 71,85-86
pKi 7.8 – 9.6 (Ki 1.6x10-8 – 2.8x10-10 M) [71,85-86]
M15 Rn Antagonist 8.0 – 9.2 pKi 71,85-86,96
pKi 8.0 – 9.2 (Ki 1x10-8 – 6.5x10-10 M) [71,85-86,96]
M15 Mm Antagonist 8.2 – 8.6 pKi 70,98
pKi 8.2 – 8.6 (Ki 6.3x10-9 – 2.24x10-9 M) [70,98]
M40 Rn Antagonist 7.9 – 8.4 pKi 71,85-86
pKi 7.9 – 8.4 (Ki 1.13x10-8 – 4x10-9 M) [71,85-86]
M871 Hs Antagonist 6.4 pKi 88
pKi 6.4 (Ki 4.2x10-7 M) [88]
Sch 202596 Hs Antagonist 5.8 pKi 16
pKi 5.8 (Ki 1.7x10-6 M) [16]
GalR3ant Hs Antagonist <5.0 pKi 52
pKi <5.0 (Ki >1x10-5 M) [52]
M35 Rn Antagonist 11.0 pIC50 33,91
pIC50 11.0 (IC50 1x10-11 M) [33,91]
M35 Hs Antagonist 10.4 pIC50 91
pIC50 10.4 (IC50 4x10-11 M) [91]
M40 Hs Antagonist 9.3 pIC50 91
pIC50 9.3 (IC50 5x10-10 M) [91]
M40 Rn Antagonist 9.1 pIC50 33,91
pIC50 9.1 (IC50 9x10-10 M) [33,91]
M15 Rn Antagonist 9.0 pIC50 33,91
pIC50 9.0 (IC50 1x10-9 M) [33,91]
M15 Hs Antagonist 8.8 – 8.8 pIC50 22,91
pIC50 8.8 – 8.8 (IC50 1.5x10-9 – 1.46x10-9 M) [22,91]
C7 Hs Antagonist 8.2 – 9.0 pIC50 22,91
pIC50 8.2 – 9.0 (IC50 6.3x10-9 – 1.04x10-9 M) [22,91]
C7 Rn Antagonist 8.3 pIC50 33,91
pIC50 8.3 (IC50 4.9x10-9 M) [33,91]
dithiipin-1,1,4,4-tetroxide analogue 7 Hs Antagonist 6.7 pIC50 84
pIC50 6.7 (IC50 1.9x10-7 M) [84]
2,3-dihydro-1,4-dithiin-1,1,4,4-tetroxide Hs Antagonist 5.6 pIC50 84
pIC50 5.6 (IC50 2.7x10-6 M) [84]
View species-specific antagonist tables
Antagonist Comments
In [84] a series of compounds was tested. The IC50 values for two of the analogues at the human GAL1 receptor are reported in the table above. Analogue 7 is reported as the first non-peptidic sub-micromolar antagonist to human GAL1 receptor.
GalR3ant displays receptor selectivity for GAL3 receptor, with low affinity for GAL1 and GAL2 receptors [53].
[24] has radioligand binding data for both membrane and whole-cell based assays.
Primary Transduction Mechanisms
Transducer Effector/Response
Gi/Go family Adenylate cyclase inhibition
Potassium channel
Comments:  Stimulation of GAL1 receptor transfected cells inhibits forskolin-stimulated cAMP production in a pertussis toxin-sensitive manner [24,29,71,85]. GAL1 receptor activation opens G-protein-regulated inwardly rectifying K+ (GIRK) channels [86] and stimulates pertussis-toxin sensitive MAPK activity, which is inhibited by expression of the C-terminus of β-adrenergic receptor kinase (which specifically inhibits G-βγ signalling). This demonstrates that GAL1 receptor couples to a Gi/βγ signalling pathway to mediate MAPK activation [95] (reviewed in [45]).
References:  7,29,71,86
Tissue Distribution
GAL1 receptor expressed in human gastrointestinal tract from oesophagus to rectum: largest amount in duodenum and least amount in gastric fundus.
Species:  Human
Technique:  RT-PCR.
References:  51
Bowes melanoma cell line > small intestine and fetal brain. None in colon adenocarcinoma cell line (HT29) or human liver.
Species:  Human
Technique:  Northern blot.
References:  29
Low expression. In periphery, most abundant in heart, small intestine, prostate, testes; in central nervous system, most abundant in cerebral cortex, amygdala, substantia nigra (not detectable in cerebellum).
Species:  Human
Technique:  RT-PCR.
References:  91
High: nucleus of the lateral olfactory tract, temporal pole of CA1 region of hippocampus.Moderate: piriform cortex, lateral septum, nucleus accumbens shell, ventral pallidum, bed nucleus stria terminalis, substantia innominata, amygdalopiriform region; anterior portion of paraventricular thalamic nucleus, centromedial nucleus, paracentral nucleus, rhomboid nucleus, subparafascicular nucleus; medial preoptic area, medial preoptic nucleus, lateral hypothalamus, perifornical region, hypothalamic paraventricular nucleus (lateral and medial zones of posterior magnocellular subdivision); central grey area of mesencephalon; external subdivision of lateral parabrachial nucleus, pontine reticular nucleus, reticulotegmental nucleus, gigantocellular, paramedian, lateral reticular nuclei, raphe magnus; dorsal horn at all levels of spinal cord (mostly laminae 1 and 2, and a few at laminae 3 and 4). Weak: glomerular and internal granular layers of the olfactory bulb, anterior olfactory nucleus, dorsal endopiriform nucleus, horizontal nucleus of the diagonal band, posterior ventral medial amygdaloid nucleus, amygdalohippocampal area; posterior intralaminar nucleus, caudal zona incerta; anterior hypothalamic area, several hypothalamic nuclei (dorsomedial, ventromedial, supraoptic, dorsal premammillary, medial mammillary, supramammillary); anterior and olivary pretectal nuclei, intermediate gray of the superior colliculus, rostral periolivary nucleus, dorsal raphe, paramedian raphe; locus coeruleus, subcoeruleus, A5 region of the pons, dorsal tegmental nucleus, medial aspect of solitary tract nucleus, dorsal vagal complex. None (or very little); sensorimotor cortex, dorsal hippocampus, cerebellum.
Species:  Rat
Technique:  In situ hybridisation.
References:  28
GAL1 receptor mRNA in > 20% of all lumbar 4 and 5 dorsal root ganglion neuron profiles, mainly small and medium sized neurons; almost all expressed CGRP mRNA. GAL1 receptor mRNA levels were transiently down-regulated by inflammation and more strongly by peripheral nerve injury
Species:  Rat
Technique:  In situ hybridisation.
References:  104
Brain; spinal cord >> small intestine. No expression in anterior pituitary, spleen, stomach, large intestine, adrenal gland, liver, lung, kidney, heart