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heterogeneous nuclear ribonucleoprotein A1
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These cells, called also delta-cells, are found in the stomach and pancreas in close proximity of G cells (Bordi et al, 2000). These cells make up approximately 2-5 % of all endocrine cells in the rat oxyntic gland and approximately 22 % in the human gland (Simonsson et al, 1988).
Cells located predominantly in the pylorus resemble pancreatic D cells (Pearse et al, 1970). Pyloric D cells have been called D1 cells by some investigators (Capella et al, 1971; Solcia et al, 1975, 1976) because their granules are smaller (0.1-0.2 micrometer) than the granules (0.2-0.4 micrometers) usually observed in gastric D cells in other species (Solcia et al, 1975, 1976). D cells can synthesize histamine at low levels and thus can be classified as APUD cells (Rubin and Schwartz, 1984).
D cells produce somatostatin and are also being referred to as alpha-1 cells or, because they produce the hormone somatostatin, as somatostatin cells or SRIF cells (SRIF being an old designation for somatostatin).
D cells and G cells are derived from a common multihormonal precursor cell (Larsson, 2000). Choi et al (2008) have implicated the homeodomain transcription factor Nkx6.3 as a selective regulator of the development of G cells and D cells, which are believed to derive from a common progenitor cell. Lee et al (2002) have shown that the bHLH transcription factor Neurogenin 3 is essential for the proper specification of gastric enteroendocrine cells and the maintenance of gastric epithelial cell identity. Knock-out mice lacking this transcription factor are characterized by the absence of D cells. These mice also lack A-cells, and gastrin secreting G cells in the epithelium of the glandular stomach. The number of serotonin-expressing enterochromaffin cells is decreased dramatically.
The density of these cells in the stomach depends on several physiological factors and is influenced by hormones and, for example, by the availability of food, surrounding pH values (Arnold et al, 1992).
D cells are important because somatostatin is the primary inhibitor of acid secretion stimulated by gastrin released by gastric G cells (Patel, 1999). Somatostatin secretion by these cells can be induced by TNF-alpha and IL8 (Beales et al, 1997), as well as IL4 (Zavros et al, 2004), whereas it is inhibited directly by IFN-gamma (Zavros et al, 2004).
D cells have been shown to possess the capacity to express the proteins listed below. Please note the following general observations, which practically apply to all cell types: 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.
¥ Activin A (EDF, erythroid differentiation factor, FRP, Follicle stimulating hormone releasing protein, Restrictin-P, WEHI-MIF, WEHI mesoderm inducing factor, activin-beta-A, INHA, inhibin-alpha, Inhibin-beta-A, INHBA ) (Yasuda H, 1993)
¥ Activin-beta B La Rosa S, 2004
¥¥ ALK7 (Activin receptor-like kinase 7, ACVR1C, activin A receptor type IC) (Bertolino et al, 2008)
¥ amylin (islet amyloid polypeptide, IAP, insulinoma amyloid polypeptide, IAPP, diabetes-associated peptide, DAP) (Mulder H, 1996; Mulder et al, 1993; Tingstedt JE, 1999; Lukinius A et al, 1996, 1997; Saruc et al, 2010)
¥ Beta-2-Microglobulin (Beta-2-M, B2M, Ly-m11, BDGF-2, bone-derived growth factor-2, CRG-8, cytokine responsive gene-8, Thymotaxin, GIP-2, granulocyte inhibitory protein, AURA54, augmented in rheumatoid arthritis 54) (Lautenschlager I, 1989)
¥¥ Bombesin (BN, BBS, GRP, Gastrin releasing peptide, gastrin releasing hormone, Mammalian Bombesin, BLP, Bombesin-like peptide) receptors (Schaffer et al, 1997)
¥ Calcitonin gene-related peptide (CALC1, CALCA, Calcitonin gene-related polypeptide-alpha, alpha-CGRP, CGRP-alpha, beta-CGRP, CGRP-beta, CGRP1, Calcitonin gene-related peptide-1, CGRP2, Calcitonin gene-related peptide-2, CALCB, Calcitonin gene-related polypeptide-beta, CALC2) (Sternini C, 1992)
¥¥ Calcitonin gene-related peptide (CALC1, CALCA, Calcitonin gene-related polypeptide-alpha, alpha-CGRP, CGRP-alpha, beta-CGRP, CGRP-beta, CGRP1, Calcitonin gene-related peptide-1, CGRP2, Calcitonin gene-related peptide-2, CALCB, Calcitonin gene-related polypeptide-beta, CALC2) receptors (Ren J et al, 1998; Kawashima et al 2002)
¥ CD54 (7F7, BB2, human rhinovirus receptor, ICAM-1, intercellular adhesion molecule 1, IFN-gamma regulated human melanoma-associated antigen, Ly47, MALA-2, Me14-D12, P3.58) (Vives M et al, 1991)
¥¥ cholecystokinin (CCK, pancreozymin) receptors (Schaffer et al, 1997; Beales et al, 1997; Dickinson CJ, 1996; Reuben M, 1994; Schmitz et al, 2001; Morisset et al, 2000)
¥ chromogranin A (CGA, CHGA, granin A, Parathyroid secretory protein-1, Parathyroid secretory protein, PSP, PSP1, pituitary secretory protein-1, Secretory protein-1, SG-I, catestatin) (Norln P et al, 2001; Cetin Y and Grube D, 1990; Portela-Gomes GM and Stridsberg M, 2001)
¥ chromogranin B (CGB, CHGB, Secretogranin-1, SCG1) (Portela-Gomes GM and Stridsberg M, 2002)
¥ Collectin-11 (COLEC11, CL-11, collectin kidney protein 1, CL-K1) (Motomura et al, 2008)
¥¥ galanin (GAL, GALN, GLNN) receptors (Fehmann HC, 1995)
¥¥ gastrin (gastrin-17, Gastrin component 3, little Gastrin, Cholecystokinin B) receptors (Helander et al, 1997)
¥¥ Glucagon (GCG, glucagon-29, pancreatic-type glucagon) receptors (Kieffer TJ, 1996)
¥¥ glucagon-like peptide-1 (GLP-1, 7-37, glucagon-like peptide 1 (7-37), GLP-1 (1-37), insulinotropin) receptors (Fehmann HC, 1995)
¥ GPR30 (G-protein-coupled receptor-30; CMKRL2; chemokine receptor-like 2; CEPR; constitutively/commonly expressed peptide-like receptor; LyGPR; lymphocyte-derived G-protein-coupled receptor; FEG1; flow induced endothelial G-protein-coupled receptor gene-1, GPER, G-protein-coupled estrogen receptor, GPER1, G-protein-coupled estrogen receptor1, DRY12, GPCR-Br) (Balhuizen A et al, 2010; Kumar et al, 2011)
¥¥ IFN-gamma (IFNG, interferon-gamma, Gamma-Interferon, gIFN) receptors (IFNGR1, IFN-gamma receptor-1, IFN-gamma-R1, interferon-gamma receptor 1, immune interferon receptor 1, AVP type 2, antiviral protein type 2, CD119, IFNGR2, interferon-gamma receptor 2, IFN-gamma receptor 2, IFN-gamma-R2, IFGR2, IFNGT1, interferon-gamma transducer 1) (Vives M et al, 1991)
¥ 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, Mecasermin) (Hansson et al, 1988, 1989)
¥¥ 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, Mecasermin) receptors (IGF1R, CD221, JTK13) (Schweiger et al, 2005)
¥ IL1-beta (IL1B, interleukin-1-beta, IL1F2, Interleukin-1 family member 2, Catabolin, H1, Hematopoietin-1, IFN-beta inducing factor, Interleukin-beta, OAF, osteoclast activating factor) (Reddy S et al, 2001)
¥¥ IL4 (Interleukin-4, BCDF-epsilon, B-cell differentiation factor-epsilon, BCDF-gamma, B-cell differentiation factor-gamma, BCGF-gamma, B-cell growth factor-gamma, BCGF-1, B-cell growth factor-1, Binetrakin, BSF-1, B-cell stimulating factor-1, BSF-p1, B-cell stimulating factor p1, EL4-BCGF, EL4 B-cell growth factor, HCGF, Hodgkin's cell growth factor, IgE-EF, IgE enhancing factor, IgG1-enhancing factor, IgG1-induction factor, LMW-BCGF, low molecular weight B-cell growth factor, MaGEF, Mast cell growth enhancing factor, MCGF-2, mast cell growth factor-2, MFF, macrophage fusion factor, Pitrakinra, TCGF-2, T-cell growth factor-2) receptors (IL4RA, IL4R-alpha, interleukin-4 receptor-alpha, IL4R, interleukin-4 receptor, CD124) (Zavros et al, 2003)
¥¥ IL8 (interleukin-8, SCYB8, 3-10C, 9E3, ANAP, anionic neutrophil-activating peptide, Chemotaxin, CEF-4, CT/IL8, CXCL8, CXC chemokine ligand 8, chemokine (C-X-C motif) ligand 8, EDNAP, endothelial-derived neutrophil-activating peptide, EMF-1, embryo fibroblast protein 1, Emoctakin, ENAP, Endothelial cell neutrophil-activating peptide, FDNAP, Fibroblast-derived neutrophil-activating peptide, FINAP, fibroblast-derived neutrophil-activating protein, GCF, granulocyte chemotactic factor, GCP, granulocyte chemotactic peptide, LAI, leukocyte adhesion inhibitor, LCF, lymphocyte chemotactic factors, LDNAP, leukocyte-derived neutrophil-activating peptide, leukocyte inhibitory factor, LUCT, lung carcinoma-derived chemotaxin, LYNAP, lymphocyte-derived neutrophil-activating peptide, MDNAP, monocyte-derived neutrophil-activating peptide, MDNCF, monocyte-derived neutrophil chemotactic factor, MOC, monocyte-derived chemotaxin, MONAP, monocyte-derived neutrophil-activating peptide, NAF, neutrophil-activating factor, NAP-1, neutrophil-activating protein-1, NCF, neutrophil chemotactic factor, NCP, neutrophil chemotactic protein, PLF, psoriatic leukotactic factor, TCF, T-cell chemotactic factor, TSG-1, Tumor necrosis factor-stimulated gene sequence-1) receptors (IL8RA, IL8R-alpha, interleukin-8 receptor-alpha; interleukin-8 receptor type A, IL8R1, IL8 receptor-1, interleukin-8 receptor type 1, CMKAR1, chemokine-alpha receptor 1, CXCR1, CD181, IL8RB, IL8R-beta, interleukin-8 receptor-beta; interleukin-8 receptor type B, IL8R2, IL8 receptor-2, interleukin-8 receptor type 2, CMKAR2, chemokine-alpha receptor 2, CXCR2, CD182) (Beales et al, 1997)
¥ LCRF (luminal CCK-releasing factor, luminal cholecystokinin releasing factor, diazepam binding inhibitor, DBI, endozepine, acyl-CoA-binding protein, ACBP, cholecystokinin releasing peptide, CCK-RP) (Ostenson et al, 1991)
¥ leptin (ob, ob/ob, obese protein, obesity factor, Metreleptin) (Vickers MH et al, 2001)
¥¥ Neuromedin B (NMB, ranatensin) receptors (Schaffer et al, 1997)
¥¥ neuromedin C (NMC, GRP-10, gastrin releasing peptide, gastrin releasing hormone, Mammalian Bombesin, BLP, Bombesin-like peptide) receptors (Schaffer et al, 1997)
¥ neuropeptide W (NPW, PPNPW, Neuropeptide W-23, NPW23, Neuropeptide W-30, NPW30, L8, L8C, GPCR8 ligand, GPCR8 ligand with c-terminal extension, PPL8, prepro-G protein-coupled receptor 8 ligand) (Rucinski et al, 2007)
¥ neuropeptide Y (NPY, Y Neuropeptide) (Myrsn U et al, 1995; Myrsn U and Sundler F, 1995)
¥ neurosecretory protein VGF [VGF, neuro-endocrine specific protein VGF, Vgf8a, TLQP-62, TLQP-21, NERP-1, Neuroendocrine regulatory peptide-1, NERP-2, Neuroendocrine regulatory peptide-2] (Brancia et al, 2010)
¥ osteopontin (OPN, OP, Osp, 2ar, 44 kDa bone phosphoprotein, 66 kDa bone phosphoprotein, bone sialoprotein, bone sialoprotein-1, BSP, BSP1, BSPI, Calcium oxalate crystal growth inhibitor protein, Eta-1, early T-lymphocyte activation protein 1, Nephropontin, Spp-1, Secreted Phosphoprotein-1, transformation-related phosphoprotein, tumor-secreted phosphoprotein, urinary stone protein, Uropontin) (Fierabracci A, 1999)
¥¥ PACAP (pituitary adenylate cyclase-activating polypeptide, ADCYAP1, adenylate cyclase-activating polypeptide 1, PACAP(1-27), PACAP27, PACAP(1-38), PACAP38) receptors (Zeng N et al, 1999)
¥ peptide YY (PYY, peptide tyrosine tyrosine) Bttcher G et al, 1994; Upchurch BH et al, 1994)
¥ RAMP-1 (receptor activity-modifying protein-1) (Kawashima K et al, 2002)
¥ Secretogranin-3 (SCG3, SgIII) (Sakai Y, 2004)
¥ Secretoneurin (SN, Chromogranin C, CHGC, SCG2, SgII, Secretogranin-2, gonadotrope polypeptide, GP-87, TSP86/84, tyrosine-sulfated protein 86/84 kDa) (Schrmann G et al, 1995; Rindi et al, 1986)
¥ somatostatin (SST, Somatotropin release inhibiting hormone, SIH, Somatotropin release inhibiting factor, SRIF, growth hormone-inhibiting hormone, GIH, growth hormone release inhibitory hormone, GH-RIH) (Zavros et al, 2003; Zeng N et al, 1999; Beales IL and Calam J, 1998; Schaffer et al, 1997; Beales et al, 1997; Dickinson CJ, 1996)
¥¥ 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 (Amherdt M et al, 1987; Ludvigsen E et al, 2004, 2005)
¥¥ substance P (Neurokinin-1, NK1, NKA, Neurokinin A, Tac1, tachykinin-1) receptors (Kwok et al, 1988)
¥¥ 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 (CD120a, TNFRSF1A, TNF receptor superfamily member 1A, FPF, TNFAR, TNFR, TNFR1, TNFR55, TNFR60, TNFR-alpha, TNF receptor 1; CD120b, TNFRSF1B, TNF receptor superfamily member 1B, p75TNFR, TNFR2, TNFR75, TNFR80, TNFR-beta) (Vives M et al, 1991; Beales IL and Calam J, 1999; Beales et al, 1997)
¥ uroguanylin (UGN, guanylate cyclase activator 2A, GUCA2B, GCAP2, guanylate cyclase activator 2, Guanylyl cyclase-activating protein 2, guanylate cyclase C activating peptide 2) (Mgert HJ et al, 1998)
¥¥ VIP (vasoactive intestinal peptide) receptors (Zeng N et al, 1999)
For other related/relevant entries see also: Cell types.
Copyright © 2012 by H IBELGAUFTS. All rights reserved.
ENTRY LAST MODIFIED: July 2012
See REFERENCES for entry D cells.
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