Cytokines & Cells Online Pathfinder Encyclopaedia
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[interstitial cells of Leydig] (a frequently misspelled form is Leyding cells) These cells are morphologically characteristic cells of the testes first described by, and named after, Franz von Leydig. The term interstitial cells is used as a synonym (hence, for example, the term Interstitial cell stimulating hormone for luteinizing hormone, which stimulates Leydig cells). Leydig cells, together with Sertoli cells, play an important role in the d evelopment and maintenance of the male phenotype and establishment of fertility (O'Shaugnessy et al, 2009).
It is thought that Leydig cell precursors migrate to the gonad from the mesonephros. During normal development of mammalian testis, 'fetal' Leydig cells appear shortly after testis differentiation in utero. An 'adult' population arises after birth (Mendis-Handagama and Ariyaratne, 2001). These two types are not two different cell types as had been suggested by earlier morphological studies (Roosen-Runge and Anderson, 1959; Lording and De Kretser, 1972). Davidoff et al (2004) have concluded from their studies that the cells of human testicular blood vessels, namely vascular smooth muscle cells and pericytes, are the progenitors of testosterone-producing Leydig cells. The undifferentiated stem Leydig cells postulated to be present in testes before the onset of precursor cell differentiation have been characterized by Ge et al, (2006).
The initial stimulus for adult Leydig cell differentiation still is not known. It may require thyroid stimulating hormone (Mendis-Handagama et al, 1998; Teerds et al, 1998), and factors produced by Sertoli cells such as Dhh and PDGF-A (Bitgood et al, 1996; Gnessi et al, 2000). Dhh signaling through its receptor, Patched-1, is involved in the early development of Leydig cells. Dhh does not appear to influence cell migration from the mesonephros to the gonad and the proliferation or survival of Leydig cell precursors but may upregulate expression of Scc in Leydig precursor cells (Yao et al, 2002). Scc is the target of the steroidogenic factor-1. Upregulation of these factors, together with other factors may trigger the differentiation of precursors into Leydig cells.
The fetal population of Leydig cells does not appear to require gonadotrophins during fetal development (O'Shaughnessy et al, 1998; Baker et al, 2001). Instead, the study of knock-out mice lacking either circulating luteninizing hormone or the hormone receptor, demonstrates that morphological and functional maturation as well as proliferation of adult Leydig cells critically depend on the presence of this hormone (Zhang et al, 2001; Lei et al, 2001).
The two populations of Leydig cells can be distinguished by a variety of markers. Scc (P450 side-chain cleavage enzyme), is a marker for fetal Leydig cells. In the mouse, adult but not fetal Leydig cells express mRNA encoding 3-beta-hydroxysteroid dehydrogenase type 6, 17-beta-hydroxysteroid dehydrogenase type 3, 11-beta-hydroxysteroid dehydrogenase type 1, prostaglandin D-synthetase, estrogen sulfotransferase, and vascular cell adhesion molecule 1 (VCAM1) (O'Shaughnessy et al, 2000, 2002; Baker et al, 1999, Baker and O'Shaughnessy, 2001; Song et al, 1997; Mendis-Handagama et al, 2001; Phillips et al, 1989). Some genes are highly expressed in the fetal Leydig cell population (renin, thrombospondin 2) but continue to be expressed at low levels in the adult testis (Perera et al, 2001, O'Shaughnessy et al, 2002).
A number of factors are known to influence the development and functional activities of Leydig cells. Sertoli cells are necessary for the survival and function of Leydig cells (Russell et al, 2001). Rat Sertoli cells have been shown to secrete a Sertoli cell factor, which affects steroidogenesis in Leydig cells, together with some biochemically uncharacterized factors that affect Leydig cell functions.
Leydig cell functions are subject to a number of factors (Haider, 2004). EGF and IGF-1 stimulate androgen production in Leydig cells. IL1 and LCI are potent Leydig cell inhibitors. Seminiferous growth factor has been shown to be a potent mitogen for Leydig cells. Part of this less characterized activity may be due to IL1. SIP [Steroidogenesis-inducing protein], from human ovarian follicular fluid, affects steroidogenesis and is a potent growth factor for Leydig cells. TGF-beta inhibits the synthesis of testosterone in Leydig cells.
The following list of mediators expressed and/or secreted by Leydig cells makes no distinction between different species. These factors may have autocrine, paracrine or endocrine functions, affecting many other cell types. Expression of most of these mediators is subject to modulation by cytokines, growth factors, and hormones.
Leydig cells have been shown to possess the capacity to express the proteins listed below. Please note the following general observation: expression may be influenced by tissue localization, may occur only in discrete subpopulations of cells, may vary between established cell lines, primary cells, embryonic cells, mature cells, fully differentiated cells, activated cells, non-activated cells or growth conditions (confluent vs. sparse cultures), may be influenced by various disease states (including cancer environment), and may differ between species.
Note also: expression profile information lists entities only for which there is an entry in COPE or one of its subdictionaries.
The meaning of • and •• is as follows: • factor/protein is expressed; •• receptor (or, in some instances, binding sites) for this factor/protein is expressed. For further explanations concerning format, "hidden" information, and/or ambiguities see my remarks in the entry cell types.
• ACTH (adrenocorticotropic hormone, corticotropin, POMC, proopiomelanocortin) (Lefebvre et al, 2003)
•• ACTH (adrenocorticotropic hormone, corticotropin, POMC, proopiomelanocortin) receptors (melanocortin receptors) (O'Shaughnessy et al, 2003)
•• Activin A (EDF, erythroid differentiation factor, FRP, Follicle stimulating hormone releasing protein, Restrictin-P, WEHI-MIF, WEHI mesoderm inducing factor, INHA, inhibin-alpha, Inhibin-beta-A, INHBA) receptors (Cameron et al, 1994)
• Activin beta-C (Gold et al, 2004)
• Activins (Anderson et al, 2002; Ying et al, 1995)
• ADAM21 (disintegrin and metalloprotease domain 21, ADAM31, disintegrin and metalloprotease domain 31) (Liu and Smith, 2000)
•• Alpha-2-Macroglobulin (Alpha-2MG, Alpha-2M, A2M) receptors (CD91) (Moestrup et al, 1992)
• ANF (Atrial natriuretic factor, natriuretic peptide, atrionatriuretic factor, Alpha-ANP, Atrial natriuretic polypeptide, ANP, Atrial polypeptide, Atriopeptin, cardionatrin, cardionatrin I, NPPA) receptors (Pandey et al, 1986; Kumar et al, 1997; Schumacher et al, 1992)
•• angiotensin-2 receptors (Khanum and Dufau, 1988)
• bFGF (basic fibroblast growth factor, FGF-2, Fibroblast growth factor-2, FGF-beta, Fibroblast growth factor-beta, HBGF-2, heparin binding growth factor-2) (Mullaney and Skinner, 1992; Ge et al, 2005)
• Bombesin (BN, BSS, GRP, gastrin releasing peptide, gastrin releasing hormone, Mammalian Bombesin, BLP, Bombesin-like peptide) (Gnessi et al, 1989)
•• bradykinin (BK, BDK, BKN, Bradykinin(1-9), BK[1-9], Alpha-2-thiol proteinase inhibitor, Fitzgerald factor, Flaujeac factor, HMWK, HMWKa, High molecular weight kininogen, HMWK-kallikrein factor, KNG, KNG1, Kininogen, Kininogen-1, LMWK, Low molecular weight kininogen, Williams factor, Williams-Fitzgerald-Flaujeac factor) receptors (Vinggaard and Hansen, 1993)
• cathepsin L (CTSL, CAL, CATL, MEP, major excreted protein, IL8 converting enzyme, CP-2, cycling protein-2, PTP, prohormone thiol convertase) (Mathur et al, 1997)
• CCL2 (CC chemokine ligand 2, chemokine (C-C motif) ligand 2, GDCF, Glioma-derived monocyte chemotactic factor-2, GDCF-2, HC11, JE, LDCF, MCAF, MCP, monocyte chemoattractant protein, MCP-1, monocyte chemoattractant protein-1, SMC-CF, smooth muscle cell chemotactic factor, TDCF, tumor-derived chemotactic factors, TSG-8, tumor necrosis factor-stimulated gene sequence-8, SCYA2) (Gerdprasert et al, 2002; Le Goffic et al, 2002; Aubry et al, 2000)
• CCL5 (CC chemokine ligand 5, chemokine (C-C motif) ligand 5, RANTES, EoCP-1, Eosinophil chemotactic polypeptide-1, SIS-delta, TCP228, T-cell-specific protein p228, SCYA5) (Le Goffic et al, 2002)
• CD56 (5.1.H11, BASCA, brain-associated small cell lung cancer antigen, Leu19, MSK39, NCAM, neural cell adhesion molecule, NCAM1, neural cell adhesion molecule-1, NKH-1, PSA-NCAM, polysialylated NCAM, E-NCAM, embryonic NCAM, embryonic neural cell adhesion molecule) (Mayerhofer et al, 1996)
• CD99 (12E7, MIC2, MIC2X, MIC2Y, E2 antigen, CD99R, O13, hec2, HBA-71) (Fellinger et al, 1991)
• CD106 (alpha-4-beta-1 ligand, INCAM-110, inducible cellular adhesion molecule 110 kDa, VCAM-1, vascular cellular adhesion molecule 1) (Verajakorva et al, 2002)
• chromogranin A (CGA, CHGA, granin A, Parathyroid secretory protein-1, Parathyroid secretory protein, PSP, PSP1, pituitary secretory protein-1, Secretory protein-1, SG-I) (Middendorff et al, 1993)
• chromogranin B (CGB, CHGB, Secretogranin-1, SCG1) (Middendorff et al, 1993)
• Clusterin (CLU, dimeric acidic glycoprotein, DAG protein, testosterone repressed prostate message-2, TRPM-2, sulfated glycoprotein-2, SGP-2, complement-associated protein SP-40,40, SP-40,40, complement lysis inhibitor, CLI, Apolipoprotein J) (Frasoldati et al, 1995)
• Corticotropin releasing hormone (CRH, CRF, Corticotropin releasing factor, Corticoliberin) (Ghizzoni et al, 1997; Tinajero et al, 1992)
•• Corticotropin releasing hormone (CRH, CRF, Corticotropin releasing factor, Corticoliberin) receptors (Ghizzoni et al, 1997)
• Cryptdins (Grandjean et al, 1997)
• C-type natriuretic peptide (CNP, ANP-C, NPPC) (Middendorff et al, 1996)
• CX3CL1 (fractalkine, FKN, FK, CX3C membrane-anchored chemokine, C3Xkine, neurotactin, NTT, NTN, ABCD-3, SCYD1) (Habasque et al, 2003)
• CXCL1 (CXC chemokine ligand 1, chemokine (C-X-C motif) ligand 1, SCYB1, CINC-1, Cytokine induced neutrophil chemoattractant-1, fsp, fibroblast secretory protein, GRO1, GRO-alpha, KC, MGSA, melanoma growth stimulatory activity, MGSA-alpha, melanoma growth stimulatory activity-alpha, NAP-3, neutrophil-activating protein-3, N51) (Le Goffic et al, 2002)
• CXCL10 (CXC chemokine ligand 10, chemokine (C-X-C motif) ligand 10, IP-10, INP-10, Interferon-inducible protein-10, IFN-inducible protein 10, immune protein-10; 10 kDa interferon-inducible protein, gamma-IP-10, CRG-2, cytokine responsive gene-2, C7, Ifi10, interferon-inducible-10, mob-1, SCYB10) (Le Goffic et al, 2002, 2003; Hu et al, 1998)
• EGF (epidermal growth factor, EGF-URO, HMGF, human milk growth factor, PGF, prostatic growth factor, beta-Urogastrone, URO, URG, Urogastrone, tooth-lid factor) (Yan et al, 1998; Zhang et al, 1997)
•• EGF (epidermal growth factor, EGF-URO, HMGF, human milk growth factor, PGF, prostatic growth factor, beta-Urogastrone, URO, URG, Urogastrone, tooth-lid factor) receptors (erb, erbB1, HER1) (Manna et al, 2002; Evaul and Hammes, 2008)
•• endothelin-1 (ET-1(1-21), ET-1) receptors (Ergul et al, 1993; Romanelli et al, 1997; Belloni et al, 1996)
•• erythropoietin (Epo, Ep, ECSA, erythroid colony stimulating activity, ESF, erythropoiesis stimulating factor) receptors (Magnanti et al, 2001; Mioni et al, 1992)
• FABP (Zschiesche et al, 1995)
• ferritin (H-Ferritin, FTH1, FTH, L-Ferritin, FTL, CD-GF, carcinoma-derived growth factor) (Koeva, 2002)
• FGF receptors (Cancilla and Risbridger, 1998; Laslett et al, 1997)
• fibronectin (FN, fibronectin-1, FN1, LETS, large external transformation-sensitive protein, Fibroblast surface antigen, SF antigen, SFA, CIg, CI globulin, cold-insoluble globulin, CSP, galactoprotein A, MSF, migration stimulating factor, Z protein) (Dickson et al, 2002)
• follistatin (FST, Follicle stimulating hormone suppressing protein, FSP, Activin-binding protein) (Anderson et al, 1998)
• gas-6 (Growth arrest-specific gene-6, Growth-potentiating factor, GPF, AXLLG, axl receptor tyrosine kinase ligand, AXSF, axl stimulatory factor) (Matsubara et al, 1997)
• Gastrin releasing peptide (GRP, gastrin releasing hormone, Mammalian Bombesin) (Gnessi et al, 1989)
•• growth hormone (GH, Somatotropic hormone, Somatotropin, GH1, growth hormone 1, GHN, growth hormone normal) receptors (Mani Maran et al, 2000)
• haptoglobin (Hp) (O'Bryan et al, 1997)
• HB-EGF (Heparin binding EGF-like factor, Heparin binding EGF-like growth factor, par-2, prostate apoptosis response protein 2, DTR, Diphtheria toxin receptor) (Vanderwinden et al, 1992)
• IFN receptors (Melaine et al, 2003)
•• IFN-gamma (IFNG, interferon-gamma, Gamma-Interferon, gIFN) receptors (CD119, IFNGR1, AVP type 2, antiviral protein type 2, immune interferon receptor) (Kanzaki and Morris, 1998)
• IGF-1 (Insulin-like growth factor-1, Erythropoietic factor, mechano growth factor, MGF, ILGF1, somatomedin C, NSILA, non-suppressible insulin-like activity, Somatomedin A, Somatomedin C, sulfation factor) (Le Roy et al, 1999; Matsui and Takahashi, 2001; Sriraman et al, 2000)
•• IGF-1 (Insulin-like growth factor-1, Erythropoietic factor, mechano growth factor, MGF, ILGF1, somatomedin C, NSILA, non-suppressible insulin-like activity, Somatomedin A, Somatomedin C, sulfation factor) receptors (IGF1R, CD221) (Le Roy et al, 1999; Sriraman et al, 2000; Zhang et al, 1998)
• IGF-2 (Insulin-like growth factor-2, Growth-promoting activity for vascular endothelial cells, T3M-11-derived growth factor, ILGF2, MSA, multiplication stimulating activity, NSILA, non-suppressible insulin-like activity, Somatomedin A, sulfation factor, SGF, skeletal growth factor, HP1-SMP, HP3-SMP, Somatomedin/insulin-like growth factor-like polypeptides) (Koike and Noumura, 1995)
• IGFBP1 (Insulin-like growth factor binding protein-1, 34 k IGFBP, AFBP, amniotic fluid binding protein, Alpha-pregnancy-associated endometrial globulin, Binding protein-25, Binding protein-26, Binding protein-28, BP-34, BP-3A, endometrial protein 14, Growth hormone independent binding protein, IGF-BP25, PP12, placental protein 12, pregnancy-associated endometrial alpha-1-globulin) (Matsui and Takahashi, 2001)
• IGFBP2 (Insulin-like growth factor binding protein-2) (Lin et al, 1993; Wang et al, 1994)
• IGFBP3 (Insulin-like growth factor binding protein-3, BP-53, Growth hormone dependent binding protein, Binding protein-29) (Wang et al, 1995; Lin et al, 1993)
• IGFBP4 (Insulin-like growth factor binding protein-4, Colon cancer cell growth inhibitor, HT29-IGFBP, HT29 insulin-like growth factor binding protein) (Hasegawa et al, 1998; Lin et al, 1993)
• IL1-alpha (IL1A, interleukin-1-alpha, BAF, B-cell activating factor, BCAF, B-cell activating factor, EP, endogenous pyrogens, LAF, lymphocyte activating factor, LEM, leukocyte endogenous mediator, MCF, mononuclear cell factor, MNCF, mononuclear cell factor, MP, mitogenic protein, TRF-3, T-cell replacing factor-3, Tumor inhibitory factor-2) (Cudicini et al, 1997)
•• IL1-alpha (IL1A, interleukin-1-alpha, BAF, B-cell activating factor, BCAF, B-cell activating factor, EP, endogenous pyrogens, LAF, lymphocyte activating factor, LEM, leukocyte endogenous mediator, MCF, mononuclear cell factor, MNCF, mononuclear cell factor, MP, mitogenic protein, TRF-3, T-cell replacing factor-3, Tumor inhibitory factor-2) receptors (IL1R, IL1RA, IL1R1, IL1 receptor type 1, IL1RB, IL1R2, IL1 receptor type 2, CD121a, CD121b) (Svechnikov et al, 2003)
• IL1-beta (IL1B, interleukin-1-beta, Catabolin, H1, Hematopoietin-1, IFN-beta inducing factor, Interleukin-beta, OAF, osteoclast activating factor) (Huleihel et al, 2003; Cudicini et al, 1997)
• IL1-beta Convertase (ICE, IL1BC, IL1-beta converting enzyme, Caspase-1, CASP1) (AbuElhija et al, 2008)
• IL1ra (IL1 receptor antagonist, IL1RN, IL1 inhibitor, IRAP, IL1 receptor antagonist protein, IL1 inhibitory activity, ILS, IL1F3, IL1 family member 3) (Huleihel et al, 2003)
•• IL1 (interleukin-1) receptors (IL1R, IL1RA, IL1R1, IL1 receptor type 1, IL1RB, IL1R2, IL1 receptor type 2, CD121a, CD121b) (Sawinski and Lukaszyk, 2002; Svechnikov et al, 2001; Lin et al, 1998)
•• IL2 (Interleukin-2, BF, blastogenic factor, EDF, eosinophil differentiation factor, KHF, killer cell helper factor, LMF, lymphocyte mitogenic factor, LPF, lymphocyte proliferation factor, MAF-C I, macrophage-activating factor for cytotoxicity I, PFC-EA, plaque forming cell enhancing factor, SCIF, secondary cytotoxic T-cell inducing factor, TCGF, T-cell growth factor, TCPA, T colony-promoting activity, TDF, thymocyte differentiation factor, T-LPF, T-lymphocyte promotor factor, TMF, thymocyte mitogenic factor, T-cell maturation factor, T-cell mitogenic factor, TRF-3, T-cell replacing factor-3, TSF, thymocyte stimulating factor) receptors (CD25, TAC antigen, Ly43, OX39) (CD122, CD132) (Guo et al, 1990; Meikle et al, 1992)
•• IL3 (Interleukin-3, 20-alpha-hydroxysteroid dehydrogenase inducing factor, BPA, burst promoting activity, CFU-S, colony-forming unit spleen, CSF-2-alpha, colony stimulating factor-2-alpha, CSF-2-beta, colony stimulating factor-2-beta, ECSF, erythroid colony stimulating factor, Eo-CSF, eosinophil colony stimulating factor, HCGF, Hematopoietic cell growth factor, HP2, Hematopoietin-2, HPGF, hematopoietic cell growth factor, MCGF, mast cell growth factor, MCSA, multi-colony stimulating activity, MEG-CSF, megakaryocyte colony stimulating factor, MGF, mast cell growth factor, Mixed colony stimulating factor, Multi-CSF, Multi-colony stimulating factor, multi-HGF, multilineage hemopoietic growth factor, Multipoietin, NC cell growth factor, natural cytotoxic cell growth factor, Neutrophil-granulocyte colony stimulating factor, PSF, progenitor stimulating factor, PSH, panspecific hemopoietin, SAF, stem cell activating factor, Thy1 inducing factor) receptors (CD123) (Morikawa et al, 1996)
• IL6 (interleukin-6, 26 kDa protein, BSF-2, B-cell stimulating factor-2, CDF, CAT development factor, choline acetyltransferase development factor, Cytolytic differentiation factor for T-lymphocytes, FDGI, fibroblast-derived growth inhibitor, HGF, hybridoma growth factor, HPGF, hybridoma/plasmacytoma growth factor, HSF, hepatocyte stimulating factor, HSF-1, hepatocyte stimulating factor-1, ILHP1, Interleukin-hemopoietin-1, MGI-2A, Macrophage-granulocyte inducer-2A, Myeloma GF, myeloma growth factor, NKAF, natural killer cell activating factor, TAF, T-cell activating factor, Thymocyte growth factor, TSF, thymocyte stimulating factor) (Kontogeorgos et al, 2002; Cudicini et al, 1997; Boockfor et al, 1994)
•• IL6 (interleukin-6, 26 kDa protein, BSF-2, B-cell stimulating factor-2, CDF, CAT development factor, choline acetyltransferase development factor, Cytolytic differentiation factor for T-lymphocytes, FDGI, fibroblast-derived growth inhibitor, HGF, hybridoma growth factor, HPGF, hybridoma/plasmacytoma growth factor, HSF, hepatocyte stimulating factor, HSF-1, hepatocyte stimulating factor-1, ILHP1, Interleukin-hemopoietin-1, MGI-2A, Macrophage-granulocyte inducer-2A, Myeloma GF, myeloma growth factor, NKAF, natural killer cell activating factor, TAF, T-cell activating factor, Thymocyte growth factor, TSF, thymocyte stimulating factor) receptors (CD126) (Sawinski and Lukaszyk, 2002)
• IL10 (interleukin-10, B-TCGF, B-cell derived T-cell growth factor, CSIF, cytokine synthesis inhibitory factor, TGIF, T-cell growth inhibitory factor) (Verajankorva et al, 2001)
• IL12 (IL12A, IL12-alpha IL12B, IL12-beta, IL12-p35, IL12-p40, IL12-p70, Interleukin-12, Interleukin-12A, Interleukin-12B, CLMF, cytotoxic lymphocyte maturation factor, NKSF, natural killer cell stimulatory factor, NKSF1, natural killer cell stimulatory factor-1, NKSF2, natural killer cell stimulatory factor-2, TcMF, CTL maturation factor, TSF, T-cell stimulating factor) (Valenti et al, 1999)
• IL18 (interleukin-18, IGIF, IFN-gamma inducing factor, IL1-gamma, IL1F4, IL1 family member 4, Iboctadekin) (AbuElhija et al, 2008)
•• IL18 (interleukin-18, IGIF, IFN-gamma inducing factor, IL1-gamma, IL1F4, IL1 family member 4, Iboctadekin) receptors (AbuElhija et al, 2008)
• inhibins (Anderson et al, 2002; Jin et al, 2001)
•• insulin receptors (CD220, INSR) (Parthasarathy and Balasubramanian, 2008; Rengarajan and Balasubramanian, 2007; Julie et al, 2003; Chatelain et al, 1987; Abelé et al, 1986)
•• Leptin (ob, ob/ob, obese protein, obesity factor) receptors (OBR, CD295, db, db/db, fatty, fa, fa/fa) (Shalitin and Phillip, 2003; Caprio et al, 2003)
• LIF (ABAE cell growth-inhibitory activity; CDF, cholinergic differentiation factor, CNDF, cholinergic neuronal differentiation factor, D-Factor, differentiation stimulating factor, DIA, differentiation inhibiting activity, DIF, differentiation inducing factor, DRF, Differentiation-retarding factor, ES cell growth factor, ESCGF, embryonic stem cell growth factor, GATS, growth stimulatory activity for TS1 cells, HILDA, human interleukin for Da cells, HSF-2, hepatocyte stimulating factor-2, HSF-3, hepatocyte stimulating factor-3, Lipoprotein lipase inhibitor, M1 differentiation inducing activity, MCGEF, mast cell growth-enhancing factor, MLPLI, melanoma-derived lipoprotein lipase inhibitor, OAF, osteoclast activating factor) (Mauduit et al, 2001; Piquet-Pellorce et al, 2000; Jenab and Morris, 1998)
•• LIF (ABAE cell growth-inhibitory activity; CDF, cholinergic differentiation factor, CNDF, cholinergic neuronal differentiation factor, D-Factor, differentiation stimulating factor, DIA, differentiation inhibiting activity, DIF, differentiation inducing factor, DRF, Differentiation-retarding factor, ES cell growth factor, ESCGF, embryonic stem cell growth factor, GATS, growth stimulatory activity for TS1 cells, HILDA, human interleukin for Da cells, HSF-2, hepatocyte stimulating factor-2, HSF-3, hepatocyte stimulating factor-3, Lipoprotein lipase inhibitor, M1 differentiation inducing activity, MCGEF, mast cell growth-enhancing factor, MLPLI, melanoma-derived lipoprotein lipase inhibitor, OAF, osteoclast activating factor) receptors (CD118) (Mauduit et al, 2001; Jenab and Morris, 1998; Lin et al, 2008)
• Luteinizing hormone receptors (Evaul and Hammes, 2008)
• M-CSF (macrophage colony stimulating factor, CSF-1, colony stimulating factor-1, CSF-HU, Urinary colony stimulating factor, MGF, macrophage growth factor, MGI-1M, macrophage-granulocyte inducer, op, osteopetrotic, toothless, tl, Lanimostim) (Ryan et al, 2001)
• MIF (MMIF, macrophage migration inhibitory factor, GIF, glycosylation inhibiting factor, phenylpyruvate tautomerase, PPT, EC5.3.2.1, phenylpyruvate keto-enol isomerase) (Wennemuth et al, 2000; Meinhardt et al, 2000)
• MIS (Müllerian inhibiting substance, AMH, anti-Müllerian hormone, MIF, Müllerian inhibiting factor, Müllerian inhibiting hormone) (Ao et al, 1993; Teixeira et al, 1999)
•• MIS (Müllerian inhibiting substance, AMH, anti-Müllerian hormone, MIF, Müllerian inhibiting factor, Müllerian inhibiting hormone) receptors (Laurich et al, 2002; Sriraman et al, 2001; Clarke et al, 2001; Teixeira et al, 1999; Lee et al, 1999; Racine et al, 1998; Fynn-Thompson et al, 1999)
• Mx (myxovirus resistance 1, Mx1, MxA, IFI-78k, interferon-inducible 78 kDa protein; IFI-78) (Melaine et al, 2003)
• neuron-specific enolase (NSE, Gamma-enolase, EC4.2.1.11, Enolase-2, ENO2, phosphopyruvate hydratase) (Middendorff et al, 1993)
• neuropeptide Y (NPY, Y Neuropeptide) (Kanzaki et al, 1996)
• neurotrophin receptors (p75NTR) (Koeva et al, 1999)
• nur-77 (nuclear receptor-77, NAK1, NGFI-B, nerve growth factor-inducible B, RNR-1, regenerating liver nuclear receptor, TIS1, TPA-inducible sequence-1, tetradecanoyl phorbol acetate-inducible sequence-1) (Martin et al, 2009; Ahn et al, 2008; El-Asmar et al, 2009)
•• NGF (nerve growth factor, 7S NGF, 7S nerve growth factor, NGF-alpha, Nerve growth factor-alpha, kallikrein Klk1b4, Klk1b4, NGF-beta, NGFB, Nerve growth factor-beta, NGFG, NGF-gamma, Nerve growth factor-gamma, kallikrein Klk1b3, Klk1b3) receptors (p75NTR) (Koeva et al, 1999)
•• NGF (nerve growth factor, 7S NGF, 7S nerve growth factor, NGF-alpha, Nerve growth factor-alpha, kallikrein Klk1b4, Klk1b4, NGF-beta, NGFB, Nerve growth factor-beta, NGFG, NGF-gamma, Nerve growth factor-gamma, kallikrein Klk1b3, Klk1b3) receptors (trkA) (Koeva et al, 1999)
•• Orexin A receptors (hypocretin 1 receptors, Ox1R) (Karteris et al, 2004)
•• Orexin B receptors (hypocretin 2 receptors, Ox2R) (Karteris et al, 2004)
• Oxytocin (OT, OXT) (Nicholson and Hardy, 1992)
•• Oxytocin (OT, OXT) receptors (Koeva and Popova, 2002; Frayne and Nicholson, 1995)
•• PACAP (pituitary adenylate cyclase-activating polypeptide, ADCYAP1, adenylate cyclase-activating polypeptide 1, PACAP(1-27), PACAP27, PACAP(1-38), PACAP38) receptors (El-Gehani et al, 2000)
• PAI-2 (Plasminogen activator inhibitor-2, placental plasminogen activator inhibitor, epidermal plasminogen activator inhibitor, monocyte-derived plasminogen activator inhibitor, Monocyte Arg-serpin, PLANH2, urokinase inhibitor, PP10, placental protein 10, Serpin B2) (Odet et al, 2006)
• patched (PTC, PTC1, PTCH, patched-1, PTCH1) (Clark et al, 2000)
• PDGF (platelet-derived growth factor, PDGF-1, PDGF-2, PDGF-A, PDGF-AA, PDGF-B, PDGF-BB, FDGF, fibroblast-derived growth factor, GDGF, glioma-derived growth factor-1, GDGF-2, glioma-derived growth factor-2, GSM, Glucocorticoid-suppressible mitogenic activity, MDF, mesangial cell proliferating factor, MDGF, monocyte-derived growth factor, OBIF, osteoblastogenesis inhibitory factor, ODGF, osteosarcoma-derived growth factor, T47D factor) (Gnessi et al, 1995; Loveland et al, 1995)
•• PDGF (platelet-derived growth factor, PDGF-1, PDGF-2, PDGF-A, PDGF-AA, PDGF-B, PDGF-BB, FDGF, fibroblast-derived growth factor, GDGF, glioma-derived growth factor-1, GDGF-2, glioma-derived growth factor-2, GSM, Glucocorticoid-suppressible mitogenic activity, MDF, mesangial cell proliferating factor, MDGF, monocyte-derived growth factor, OBIF, osteoblastogenesis inhibitory factor, ODGF, osteosarcoma-derived growth factor, T47D factor) receptors (Gnessi et al, 1995; Loveland et al, 1995; Risbridger, 1993; Ge et al, 2005)
•• prolactin (PRL, lactogenic hormone, lactotropin, lactotropic hormone, mammotropin, mammotropic hormone, luteotropic hormone, LTH, luteotropin) receptors (Maran et al, 2001; Hennuy et al, 2000; Weiss-Messer et al, 1998; Hair et al, 2002; Parthasarathy and Balasubramanian , 2008)
• proopiomelanocortin (POMC, ACTH, ACTH[1-13], ACTH[18-39], ACTH[22-39], adrenocorticotropin, Alpha-Endorphin, alpha-melanocortin, alpha-melanotropin, Alpha-MSH, Beta-cell trophin, Beta-Endorphin, Beta-Lipotropin, Beta-LPH, beta-melanotropin, Beta-MSH, CLIP, corticotrophin, corticotropin, corticotropin-like intermediate peptide, Gamma-Endorphin, Gamma-Lipotropin, Gamma-LPH, Gamma-MSH, Lipotropin, melanocyte stimulating hormone-alpha, melanocyte stimulating hormone-beta, melanocyte stimulating hormone-gamma, melanotropin, POC, pro-ACTH-Endorphin, proopiocortin) (Ivell, 1994; Li et al, 1993)
• PTHrP (parathyroid hormone-like related protein, PTH-related protein, parathyroid hormone-related protein, PTHR, PTH-related peptide, Parathyroid hormone-related peptide, parathyroid hormone-like hormone, PTHLH) (Campos et al, 1991; Aklilu et al, 2000)
•• Relaxin (RLN, RLX, RLN1, relaxin 1, relaxin H1, H1 relaxin, RLXH1, RXN1, RLN2, relaxin 2, relaxin H2, H2 relaxin, RLXH2, RXN2, relaxin 1/2) receptors (Min et al, 1998)
• relaxin-like factor (RLF, Leydig cell insulin-like factor, Ley-I-L, Insulin-like-3, INSL3) (Lague and Tremblay, 2008; Bamberger et al, 1999; Kawamura et al, 2004; Kohsaka et al, 2009; Koeva et al, 2008; Balvers et al, 1998; Ivell et al, 1997; Paust et al, 2002; Pusch et al, 1996; Feng et al, 2007; Arrighi et al, 2009)
•• relaxin-like factor (RLF, Leydig cell insulin-like factor, Ley-I-L, Insulin-like-3, INSL3) receptors (Anand-Ivell RJ et al, 2006; Kohsaka et al, 2009; Koeva et al, 2008)
• RXFP1 (LGR7, leucine-rich repeat-containing G-protein-coupled receptor 7, RLNR, relaxin receptor) (Kohsaka et al, 2009
• RXFP2 (relaxin family peptide receptor 2, LGR8, leucine-rich repeat-containing G-protein-coupled receptor 8, GREAT, G-protein-coupled receptor affecting testicular descent, INSL3R, INSL3 receptor) (Anand-Ivell et al, 2006; Koeva et al, 2008; Feng et al, 2007)
• S100 (S100a, S100A1, S100-alpha, S100b, S100-beta, NEF, neurite extension factor) (Middendorff et al, 1993)
• SCF (stem cell factor, HLGF-1, hemolymphopoietic growth factor-1), KL, kit ligand, KITL, KITLG, Mast/stem cell growth factor, MGF, mast cell growth factor, SLF, steel factor) (Munsie et al, 1997; Kondoh et al, 1995)
•• SCF (stem cell factor, HLGF-1, hemolymphopoietic growth factor-1), KL, kit ligand, KITL, KITLG, Mast/stem cell growth factor, MGF, mast cell growth factor, SLF, steel factor) receptors (kit, CD117) (Yan et al, 2000; Feng et al, 1999; Tsuchida et al, 2003; Kondoh et al, 1995; Ge et al, 2005)
• Sertoli cell secreted growth factor (SCSGF) (Buch et al, 1988; Lamb et al, 1991)
• SMAD proteins (Hu et al, 2003; Clarke et al, 2001)
• somatostatin (SST, Somatotropin release inhibiting hormone, SIH, Somatotropin release inhibiting factor, SRIF, growth hormone-inhibiting hormone, GIH, growth hormone release inhibitory hormone, GH-RIH) (Goddard et al, 2001; Fombonne et al, 2003)
•• somatostatin (SST, Somatotropin release inhibiting hormone, SIH, Somatotropin release inhibiting factor, SRIF, growth hormone-inhibiting hormone, GIH, growth hormone release inhibitory hormone, GH-RIH) receptors (Fombonne et al, 2003)
• SPARC (secreted protein acidic and rich in cysteine, osteonectin, BM-40, 43K glycoprotein) (Vernon and Sage, 1989)
• stanniocalcin 1 (STC1, STC, STC50, stanniocalcin, hypocalcin, teleocalcin) (Li and Wong, 2008)
• substance P (Neurokinin-1, NK1, NKA, Neurokinin A, Tac1, tachykinin-1) (Middendorff et al, 1993; Chiwakata et al, 1991)
• Survivin (SVV, Apoptosis inhibitor 4, API4, TIAP, Thiol inhibitor of apoptosis, BIRC5, baculoviral IAP repeat-containing protein-5) (Wang et al, 2004)
• TCTP (translationally controlled tumor protein, TPT1, tumor protein translationally-controlled-1, fortilin, p23, IgE-HRF, IgE-dependent histamine releasing factor) (Guillaume et al, 2001)
• TGF-beta (transforming growth factor-beta, TGFB, B-TGF, Aqueous humor lymphocyte inhibitory activity, DIF, differentiation-inhibiting factor, EGI, epithelial cell-specific growth inhibitor; epithelial growth inhibitor, EIF, Epstein-Barr virus inducing factor, Epithelial cell growth inhibiting factor, G-TsF, glioma-derived T-cell suppressor factor, MDGF, milk-derived growth factor, MGF, milk growth factor, Polyergin, Simian BSC-1 cell growth inhibitor, SP factor, TCGF, transformed cell growth factor, TGI, tissue-derived growth inhibitor, TIF-1, tumor inducing factor-1) (Le Roy et al, 1999; Olaso et al, 1999; Caussanel et al, 1997)
•• TGF-beta (transforming growth factor-beta, TGFB, B-TGF, Aqueous humor lymphocyte inhibitory activity, DIF, differentiation-inhibiting factor, EGI, epithelial cell-specific growth inhibitor; epithelial growth inhibitor, EIF, Epstein-Barr virus inducing factor, Epithelial cell growth inhibiting factor, G-TsF, glioma-derived T-cell suppressor factor, MDGF, milk-derived growth factor, MGF, milk growth factor, Polyergin, Simian BSC-1 cell growth inhibitor, SP factor, TCGF, transformed cell growth factor, TGI, tissue-derived growth inhibitor, TIF-1, tumor inducing factor-1) receptors (Dickson et al, 2002; Goddard et al, 2000; Le Roy et al, 1999; Olaso et al, 1998; Khan et al, 1999)
• TIMP-2 (tissue inhibitor of metalloproteinases-2, TIMP metallopeptidase inhibitor 2, CSC-21K, MI, CHIAMP, Chondrocyte-derived inhibitor of angiogenesis and metalloproteinase activity) (Ge et al, 2005)
•• TNF-alpha (tumor necrosis factor-alpha, TNFSF2, TNF ligand superfamily member 2, Cachectin, CF, cytotoxic factor, CTX, cytotoxin, DIF, differentiation inducing factor, EP, endogenous pyrogens, Hemorrhagic factor, Macrophage-derived cytotoxic factor, J774-derived cytotoxic factor, MCF, macrophage cytotoxic factor, MCT, macrophage cytotoxin, MD-FGF, monocyte-derived fibroblast growth factor, PCF, peritoneal cytotoxic factor, RCF, Released cytotoxic factor) receptors (Budnik et al, 1999; Rey et al, 2000)
• TRAIL (TNF-related apoptosis inducing ligand, APO-2 ligand, APO-2L, TL2, TNF-like-2, TNF ligand superfamily member 10, TNFSF10, Ly81, CD253) (Grataroli et al, 2004)
•• TRAIL (TNF-related apoptosis inducing ligand, APO-2 ligand, APO-2L, TL2, TNF-like-2, TNF ligand superfamily member 10, TNFSF10, Ly81, CD253) receptor-2 (TRAIL receptor-2, TRAIL-R2, death receptor-5, DR5, KILLER, TRICK2, TRAIL receptor inducer of cell killing-2, TNFRSF10B, TNF receptor superfamily member 10B, CD262, Ly98) (Grataroli et al, 2002)
•• TRAIL (TNF-related apoptosis inducing ligand, APO-2 ligand, APO-2L, TL2, TNF-like-2, TNF ligand superfamily member 10, TNFSF10, Ly81, CD253) receptor-4 (TRAIL receptor-4, TRAIL-R4, DcR-2, Decoy receptor-2, TRUNDD, TRAIL receptor with a truncated death domain, TNFRSF10D, TNF receptor superfamily member 10D, CD264) (Grataroli et al, 2002, 2004)
•• urocortin (UCN, Urocortin-1, Urotensin-1) receptors (Rivier, 2008)
• Vasopressin (Arginine vasopressin, Arg-Vasopressin, AVP, ARVP, VP, Antidiuretic hormone, ADH, Adiuretin, Vasotocin, Pituitrin P, Pitressin, arginine-vasopressin-neurophysin II, vasopressin-neurophysin 2-copeptin, AVP-NpII) (Ivell et al, 1992)
•• Vasopressin (Arginine vasopressin, Arg-Vasopressin, AVP, ARVP, VP, Antidiuretic hormone, ADH, Adiuretin, Vasotocin, Pituitrin P, Pitressin, arginine-vasopressin-neurophysin II, vasopressin-neurophysin 2-copeptin, AVP-NpII) receptors (Hales and Greene, 1998; Frayne and Nicholson, 1995; Vinggaard and Hansen, 1993)
• VEGF (vascular endothelial growth factor, VEGF-A, vascular endothelial growth factor A, VEGF-1, vascular endothelial growth factor-1, VEG/PF, vascular endothelial growth factor/vascular permeability factor, GD-VEGF, glioma-derived vascular endothelial growth factor, VAS, Vasculotropin, Vascular endothelial cell proliferation factor, VPF, vascular permeability factor, FSdGF, Folliculo stellate cell-derived growth factor) (Anand et al, 2003; Ergun et al, 1997)
•• VEGF (vascular endothelial growth factor, VEGF-A, vascular endothelial growth factor A, VEGF-1, vascular endothelial growth factor-1, VEG/PF, vascular endothelial growth factor/vascular permeability factor, GD-VEGF, glioma-derived vascular endothelial growth factor, VAS, Vasculotropin, Vascular endothelial cell proliferation factor, VPF, vascular permeability factor, FSdGF, Folliculo stellate cell-derived growth factor) receptors (Ergun et al, 1997)
• vitronectin (VN, VTN, Epibolin, Serum spreading factor, SF, Membrane attack complex inhibitor, S protein, Protein X, Complement S protein, PAIBP, Plasminogen activator inhibitor-1 binding protein, Somatomedin B) (Sawada et al, 1996)
For related information see also: Cell types, Cell lines in Cytokine Research, Cell culture.
• Wnt-2b (Wnt-13, Wnt-2b1, Wnt-2b2) (Fowler et al, 2007)
•• Wnt-4 receptors (Jordan et al, 2001, 2003)
LAST MODIFIED: October 2009
See REFERENCES for entry Leydig cells
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