Membrane steroid receptor

Membrane steroid receptors (mSRs), also called extranuclear steroid receptors, are a class of receptors that bind and are activated by endogenous steroids and mediate rapid, non-genomic signaling via modulation of intracellular signaling cascades.^[1]^[2]^[3]^[4] They are another means besides classical nuclear steroid hormone receptors by which steroids mediate their biological effects.^[1]^[2]^[3]^[4]

Known groups of mSRs include:

Membrane sex steroid receptors
- Membrane estrogen receptors (mERs) – e.g., GPER, ER-X, G_q-mER
- Membrane progesterone receptors (mPRs) – mPRα, mPRβ, mPRγ
- Membrane androgen receptors (mARs) – GPRC6A, ZIP9
Membrane corticosteroid receptors
- Membrane glucocorticoid receptors (mGRs)
- Membrane mineralocorticoid receptors (mMRs)

Ligand-gated ion channels such as the GABA_A receptor and the NMDA receptor bind and are modulated by neurosteroids such as dehydroepiandrosterone (DHEA), allopregnanolone, and 3α-androstanediol. Vomeronasal receptors bind and are activated by pheromones such as androstenone and androstenol.

References

1 2 Levin ER (2014). "Translating extranuclear steroid receptor signaling to clinical medicine". Horm Cancer. 5 (3): 140–5. doi:10.1007/s12672-014-0179-9. PMC 4040225. PMID 24752388.
1 2 Hammes SR, Levin ER (2011). "Minireview: Recent advances in extranuclear steroid receptor actions". Endocrinology. 152 (12): 4489–95. doi:10.1210/en.2011-1470. PMC 3858720. PMID 22028449.
1 2 Sen A, Prizant H, Hammes SR (2011). "Understanding extranuclear (nongenomic) androgen signaling: what a frog oocyte can tell us about human biology". Steroids. 76 (9): 822–8. doi:10.1016/j.steroids.2011.02.016. PMID 21354434.
1 2 Watson, C. S. (1999). "Signaling Themes Shared Between Peptide and Steroid Hormones at the Plasma Membrane". Science Signaling. 1999 (12): pe1–pe1. doi:10.1126/stke.1999.12.pe1. ISSN 1945-0877.

Cell surface receptor: G protein-coupled receptors

Class A: Rhodopsin like

Neurotransmitter

Adrenergic	α1 (A B D) α2 (A B C) β1 β2 β3

Purinergic	Adenosine (A1 A2A A2B A3) P2Y (1 2 4 5 6 8 9 10 11 12 13 14)

Serotonin	(all but 5-HT3) 5-HT1 (A B D E F) 5-HT2 (A B C) 5-HT (4 5A 6 7)

Other	Acetylcholine (M1 M2 M3 M4 M5) Dopamine (D1 D2 D3 D4 D5) Histamine (H1 H2 H3 H4) Melatonin (1A 1B 1C) TAAR (1 2 3 5 6 8 9)

Metabolites and
signaling molecules

Eicosanoid	CysLT (1 2) LTB4 (1 2) FPRL1 OXE Prostaglandin (DP (1 2), EP (1 2 3 4), FP) Prostacyclin Thromboxane

Other	Bile acid Cannabinoid (CB1 CB2, GPR (18 55 119)) EBI2 Estrogen Free fatty acid (1 2 3 4) Lactate Lysophosphatidic acid (1 2 3 4 5 6) Lysophospholipid (1 2 3 4 5 6 7 8) Niacin (1 2) Oxoglutarate PAF Sphingosine-1-phosphate (1 2 3 4 5) Succinate

Peptide

Neuropeptide	B/W (1 2) FF (1 2) S Y (1 2 4 5) Neuromedin (B U (1 2)) Neurotensin (1 2)

Other	Anaphylatoxin (C3a C5a) Angiotensin (1 2) Apelin Bombesin (BRS3 GRPR NMBR) Bradykinin (B1 B2) Chemokine Cholecystokinin (A B) Endothelin (A B) Formyl peptide (1 2 3) FSH Galanin (1 2 3) GHB receptor Gonadotropin-releasing hormone (1 2) Ghrelin Kisspeptin Luteinizing hormone/choriogonadotropin MAS (1 1L D E F G X1 X2 X3 X4) Melanocortin (1 2 3 4 5) MCHR (1 2) Motilin Opioid (Delta Kappa Mu Nociceptin & Zeta, but not Sigma) Orexin (1 2) Oxytocin Prokineticin (1 2) Prolactin-releasing peptide Relaxin (1 2 3 4) Somatostatin (1 2 3 4 5) Tachykinin (1 2 3) Thyrotropin Thyrotropin-releasing hormone Urotensin-II Vasopressin (1A 1B 2)

Miscellaneous

Orphan	GPR (1 3 4 6 12 15 17 18 19 20 21 22 23 25 26 27 31 32 33 34 35 37 39 42 44 45 50 52 55 61 62 63 65 68 75 77 78 81 82 83 84 85 87 88 92 101 103 109A 109B 119 120 132 135 137B 139 141 142 146 148 149 150 151 152 153 160 161 162 171 173 174 176 177 182 183)

Other	Adrenomedullin Olfactory Opsin (3 4 5 1LW 1MW 1SW RGR RRH) Protease-activated (1 2 3 4) SREB (1 2 3)

Class B: Secretin like

Adhesion	ADGRG (1 2 3 4 5 6 7)

Orphan	GPR (56 64 97 98 110 111 112 113 114 115 116 123 124 125 126 128 133 143 144 155 157)

Other	Brain-specific angiogenesis inhibitor (1 2 3) Cadherin (1 2 3) Calcitonin CALCRL CD97 Corticotropin-releasing hormone (1 2) EMR (1 2 3) Glucagon (GR GIPR GLP1R GLP2R) Growth hormone releasing hormone PACAPR1 GPR Latrophilin (1 2 3 ELTD1) Methuselah-like proteins Parathyroid hormone (1 2) Secretin Vasoactive intestinal peptide (1 2)

Class C: Metabotropic glutamate / pheromone

Taste	TAS1R (1 2 3) TAS2R (1 3 4 5 7 8 9 10 13 14 16 19 20 30 31 38 39 40 41 42 43 45 46 50 60 Vomeronasal receptor)

Other	Calcium-sensing receptor GABA B (1 2) Glutamate receptor (Metabotropic glutamate (1 2 3 4 5 6 7 8)) GPRC6A GPR (156 158 179) RAIG (1 2 3 4)

Class F: Frizzled / Smoothened

Frizzled	Frizzled (1 2 3 4 5 6 7 8 9 10)

Smoothened	Smoothened

Endogenous steroids

Precursors

Corticosteroids

Glucocorticoids	3α,5α-Tetrahydrocorticosterone 5α-Dihydrocorticosterone 11-Deoxycorticosterone 11-Deoxycortisol 21-Deoxycortisol Corticosterone Cortisol Cortisone 17α-Hydroxypregnenolone 17α-Hydroxyprogesterone Pregnenolone Progesterone

Mineralocorticoids	5α-Dihydroaldosterone 11-Dehydrocorticosterone (11-oxocorticosterone, 17-deoxycortisone) 11-Deoxycortisol 11-Deoxycorticosterone 11β-Hydroxyprogesterone (21-deoxycorticosterone) 16α,18-Dihydroxy-11-deoxycorticosterone 18-Hydroxy-11-deoxycorticosterone 18-Hydroxycorticosterone Aldosterone Corticosterone Cortisol

Sex steroids

Androgens	11-Ketodihydrotestosterone 11-Ketotestosterone 7β-Hydroxyepiandrosterone 11β-Hydroxyandrostenedione 4-Androstenedione 5-Androstenediol Adrenosterone (11-ketoandrostenedione) Androsterone DHEA DHEA sulfate Dihydrotestosterone Epiandrosterone Epitestosterone 16α-Hydroxyandrostenedione 16α-Hydroxy-DHEA 16α-Hydroxy-DHEA sulfate Testosterone Metabolites: 3α-Androstanediol 3α-Androstanediol glucuronide 3β-Androstanediol 3α,5β-Androstanediol 3β,5β-Androstanediol 5β-Androstanedione 5β-Dihydrotestosterone Androstanetriols Androsterone glucuronide Dihydrotestosterone glucuronide Dihydrotestosterone sulfate Etiocholanolone Etiocholanolone glucuronide Epietiocholanolone Testosterone glucuronide Testosterone sulfate

Estrogens	Estranes: Estetrol Estradiol Estrone Estriol 16β-Epiestriol (16β-hydroxyestradiol) 17α-Epiestriol (16α-hydroxy-17α-estradiol) 16β,17α-Epiestriol (16β-hydroxy-17α-estradiol) 2-Hydroxyestradiol 2-Hydroxyestrone 2-Methoxyestradiol 4-Hydroxyestradiol 4-Hydroxyestrone 16α-Hydroxyestrone Others: 27-Hydroxycholesterol 3α-Androstanediol 3β-Androstanediol 4-Androstenedione 5-Androstenediol DHEA DHEA sulfate 7-Keto-DHEA 7α-Hydroxy-DHEA 16α-Hydroxy-DHEA Metabolites: Estradiol glucuronide Estradiol sulfate Estrone glucuronide Estrone sulfate Estriol glucuronide Estriol sulfate Lipoidal estradiol (e.g., estradiol stearate, estradiol palmitate)

Progestogens	Progesterone 16α-Hydroxyprogesterone 17α-Hydroxyprogesterone 20α-Dihydroprogesterone 5α-Dihydroprogesterone 11-Deoxycorticosterone 5α-DHDOC Metabolites: Allopregnanediol Pregnanediol Pregnanediol glucuronide Pregnanetriol

Neurosteroids

Cholestanes: 24S-Hydroxycholesterol
Cholesterol

Androstanes: 3α-Androstanediol
3α-Androstenol
7-Keto-DHEA
7α-Hydroxy-DHEA
7β-Hydroxy-DHEA
7α-Hydroxyepiandrosterone
7β-Hydroxyepiandrosterone
Androsterone
DHEA
DHEA sulfate
Etiocholanolone

Others

Others: 11α-Hydroxyprogesterone
11β-Hydroxyprogesterone

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