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{{Short description|Protein-coding gene in the species Homo sapiens}} |
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{{Infobox_gene}} |
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'''Interleukin 9''', also known as '''IL-9''', is a [[Pleiotropy|pleiotropic]] [[cytokine]] (cell signalling molecule) belonging to the group of [[interleukin]]s.<ref name="entrez">{{cite web | title = Entrez Gene: IL9 interleukin 9| url = https://www.ncbi.nlm.nih.gov/sites/entrez?Db=gene&Cmd=ShowDetailView&TermToSearch=3578}}</ref> IL-9 is produced by variety of cells like [[mast cell]]s, [[Natural killer T cell|NKT]] cells, [[T helper cell|Th2]], [[T helper 17 cell|Th17]], [[Regulatory T cell|Treg]], [[ILC2]], and Th9 cells in different amounts. Among them, [[T helper cell|Th9 cells]] are regarded as the major [[CD4+ helper cells|CD4+ T cells]] that produce IL-9.<ref name="ReferenceA">{{cite book | vauthors = Rojas-Zuleta WG, Sanchez E | title = Th9 Cells | chapter = IL-9: Function, Sources, and Detection | series = Methods in Molecular Biology | volume = 1585 | pages = 21–35 | date = 2017 | pmid = 28477184 | doi = 10.1007/978-1-4939-6877-0_2 | isbn = 978-1-4939-6876-3 | hdl = 10161/14730 }}</ref> |
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== Functions == |
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{{GNF_Protein_box |
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| image = |
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| image_source = |
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| PDB = |
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| Name = Interleukin 9 |
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| HGNCid = 6029 |
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| Symbol = IL9 |
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| AltSymbols =; HP40; IL-9; P40 |
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| OMIM = 146931 |
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| ECnumber = |
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| Homologene = 492 |
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| MGIid = 96563 |
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| GeneAtlas_image1 = PBB_GE_IL9_208193_at_tn.png |
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| Function = {{GNF_GO|id=GO:0005140 |text = interleukin-9 receptor binding}} |
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| Component = {{GNF_GO|id=GO:0005576 |text = extracellular region}} {{GNF_GO|id=GO:0005615 |text = extracellular space}} |
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| Process = {{GNF_GO|id=GO:0006954 |text = inflammatory response}} {{GNF_GO|id=GO:0006955 |text = immune response}} {{GNF_GO|id=GO:0007267 |text = cell-cell signaling}} {{GNF_GO|id=GO:0008284 |text = positive regulation of cell proliferation}} {{GNF_GO|id=GO:0045407 |text = positive regulation of interleukin-5 biosynthetic process}} |
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| Orthologs = {{GNF_Ortholog_box |
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| Hs_EntrezGene = 3578 |
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| Hs_Ensembl = ENSG00000145839 |
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| Hs_RefseqProtein = NP_000581 |
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| Hs_RefseqmRNA = NM_000590 |
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| Hs_GenLoc_db = |
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| Hs_GenLoc_chr = 5 |
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| Hs_GenLoc_start = 135255834 |
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| Hs_GenLoc_end = 135259415 |
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| Hs_Uniprot = P15248 |
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| Mm_EntrezGene = 16198 |
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| Mm_Ensembl = ENSMUSG00000021538 |
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| Mm_RefseqmRNA = NM_008373 |
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| Mm_RefseqProtein = NP_032399 |
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| Mm_GenLoc_db = |
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| Mm_GenLoc_chr = 13 |
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| Mm_GenLoc_start = 56488899 |
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| Mm_GenLoc_end = 56491868 |
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| Mm_Uniprot = P15247 |
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}} |
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}} |
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'''Interleukin 9''', also known as '''IL9''', is a [[cytokine]] (cell signalling molecule) belonging to the group of [[interleukin]]s.<ref name="entrez">{{cite web | title = Entrez Gene: IL9 interleukin 9| url = http://www.ncbi.nlm.nih.gov/sites/entrez?Db=gene&Cmd=ShowDetailView&TermToSearch=3578| accessdate = }}</ref> |
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Il-9 is a [[cytokine]] secreted by [[CD4+ helper cells]] that acts as a regulator of a variety of [[hematopoietic cell]]s.<ref name="pmid21371941">{{cite journal | vauthors = Perumal NB, Kaplan MH | title = Regulating IL9 transcription in T helper cells | journal = Trends in Immunology | volume = 32 | issue = 4 | pages = 146–50 | year = 2011 | pmid = 21371941 | pmc = 3070825 | doi = 10.1016/j.it.2011.01.006 }}</ref> This cytokine stimulates [[cell proliferation]] and prevents [[apoptosis]]. It functions through the [[interleukin-9 receptor]] (IL9R), which activates different signal transducer and activator ([[STAT protein|STAT]]) proteins namely [[STAT1]], [[STAT3]] and [[STAT5]] and thus connects this cytokine to various biological processes. The gene encoding this cytokine has been identified as a candidate gene for [[asthma]]. Genetic studies on a mouse model of asthma demonstrated that this cytokine is a determining factor in the pathogenesis of [[bronchial hyperresponsiveness]].<ref name="entrez" /> |
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{{PBB_Summary |
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| section_title = |
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| summary_text = The protein encoded by this gene is a cytokine produced by T-cells and specifically by CD4+ helper cells that acts as a regulator of a variety of hematopoietic cells. This cytokine stimulates cell proliferation and prevents apoptosis. It functions through the [[interleukin-9 receptor]] (IL9R), which activates different signal transducer and activator (STAT) proteins and thus connects this cytokine to various biological processes. The gene encoding this cytokine has been identified as a candidate gene for [[asthma]]. Genetic studies on a mouse model of asthma demonstrated that this cytokine is a determining factor in the pathogenesis of bronchial hyperresponsiveness.<ref name="entrez">{{cite web | title = Entrez Gene: IL9 interleukin 9| url = http://www.ncbi.nlm.nih.gov/sites/entrez?Db=gene&Cmd=ShowDetailView&TermToSearch=3578| accessdate = }}</ref> |
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}} |
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Interleukin-9 has also shown to inhibit [[melanoma]] growth in mice. |
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==References== |
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<ref name="pmid22772464">{{cite journal | vauthors = Purwar R, Schlapbach C, Xiao S, Kang HS, Elyaman W, Jiang X, Jetten AM, Khoury SJ, Fuhlbrigge RC, Kuchroo VK, Clark RA, Kupper TS | title = Robust tumor immunity to melanoma mediated by interleukin-9-producing T cells | journal = Nature Medicine | volume = 18 | issue = 8 | pages = 1248–53 | date = August 2012 | pmid = 22772464 | pmc = 3518666 | doi = 10.1038/nm.2856 }} |
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{{reflist}} |
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*{{lay source |template = cite web|url = https://medicalxpress.com/news/2012-07-scientists-molecule-immune-dangerous-skin.html|title = Scientists find molecule in immune system that could help treat dangerous skin cancer|date = July 8, 2012 |website = MedicalXpress }}</ref> |
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Additionally, it gives rise to the multiplication of [[Hematologic neoplasms|hematologic neoplasias]] and also [[Hodgkin's lymphoma]] in humans but IL-9 also has antitumor properties in solid tumors, for example [[melanoma]].<ref name="ReferenceA"/> |
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==Further reading== |
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{{refbegin | 2}} |
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{{PBB_Further_reading |
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| citations = |
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*{{cite journal | author=Renauld JC, Houssiau F, Louahed J, ''et al.'' |title=Interleukin-9. |journal=Adv. Immunol. |volume=54 |issue= |pages= 79-97 |year= 1993 |pmid= 8379467 |doi= }} |
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*{{cite journal | author=Knoops L, Renauld JC |title=IL-9 and its receptor: from signal transduction to tumorigenesis. |journal=Growth Factors |volume=22 |issue= 4 |pages= 207-15 |year= 2005 |pmid= 15621723 |doi= 10.1080/08977190410001720879 }} |
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*{{cite journal | author=Modi WS, Pollock DD, Mock BA, ''et al.'' |title=Regional localization of the human glutaminase (GLS) and interleukin-9 (IL9) genes by in situ hybridization. |journal=Cytogenet. Cell Genet. |volume=57 |issue= 2-3 |pages= 114-6 |year= 1991 |pmid= 1680606 |doi= }} |
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*{{cite journal | author=Kelleher K, Bean K, Clark SC, ''et al.'' |title=Human interleukin-9: genomic sequence, chromosomal location, and sequences essential for its expression in human T-cell leukemia virus (HTLV)-I-transformed human T cells. |journal=Blood |volume=77 |issue= 7 |pages= 1436-41 |year= 1991 |pmid= 1901233 |doi= }} |
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*{{cite journal | author=Holbrook ST, Ohls RK, Schibler KR, ''et al.'' |title=Effect of interleukin-9 on clonogenic maturation and cell-cycle status of fetal and adult hematopoietic progenitors. |journal=Blood |volume=77 |issue= 10 |pages= 2129-34 |year= 1991 |pmid= 1903074 |doi= }} |
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*{{cite journal | author=Merz H, Houssiau FA, Orscheschek K, ''et al.'' |title=Interleukin-9 expression in human malignant lymphomas: unique association with Hodgkin's disease and large cell anaplastic lymphoma. |journal=Blood |volume=78 |issue= 5 |pages= 1311-7 |year= 1991 |pmid= 1908723 |doi= }} |
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*{{cite journal | author=Renauld JC, Goethals A, Houssiau F, ''et al.'' |title=Human P40/IL-9. Expression in activated CD4+ T cells, genomic organization, and comparison with the mouse gene. |journal=J. Immunol. |volume=144 |issue= 11 |pages= 4235-41 |year= 1990 |pmid= 1971295 |doi= }} |
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*{{cite journal | author=Renauld JC, Goethals A, Houssiau F, ''et al.'' |title=Cloning and expression of a cDNA for the human homolog of mouse T cell and mast cell growth factor P40. |journal=Cytokine |volume=2 |issue= 1 |pages= 9-12 |year= 1991 |pmid= 2129501 |doi= }} |
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*{{cite journal | author=Yang YC, Ricciardi S, Ciarletta A, ''et al.'' |title=Expression cloning of cDNA encoding a novel human hematopoietic growth factor: human homologue of murine T-cell growth factor P40. |journal=Blood |volume=74 |issue= 6 |pages= 1880-4 |year= 1989 |pmid= 2508790 |doi= }} |
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*{{cite journal | author=Yin T, Keller SR, Quelle FW, ''et al.'' |title=Interleukin-9 induces tyrosine phosphorylation of insulin receptor substrate-1 via JAK tyrosine kinases. |journal=J. Biol. Chem. |volume=270 |issue= 35 |pages= 20497-502 |year= 1995 |pmid= 7544789 |doi= }} |
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*{{cite journal | author=Postma DS, Bleecker ER, Amelung PJ, ''et al.'' |title=Genetic susceptibility to asthma--bronchial hyperresponsiveness coinherited with a major gene for atopy. |journal=N. Engl. J. Med. |volume=333 |issue= 14 |pages= 894-900 |year= 1995 |pmid= 7666875 |doi= }} |
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*{{cite journal | author=Le Beau MM, Espinosa R, Neuman WL, ''et al.'' |title=Cytogenetic and molecular delineation of the smallest commonly deleted region of chromosome 5 in malignant myeloid diseases. |journal=Proc. Natl. Acad. Sci. U.S.A. |volume=90 |issue= 12 |pages= 5484-8 |year= 1993 |pmid= 8516290 |doi= }} |
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*{{cite journal | author=Demoulin JB, Uyttenhove C, Van Roost E, ''et al.'' |title=A single tyrosine of the interleukin-9 (IL-9) receptor is required for STAT activation, antiapoptotic activity, and growth regulation by IL-9. |journal=Mol. Cell. Biol. |volume=16 |issue= 9 |pages= 4710-6 |year= 1996 |pmid= 8756628 |doi= }} |
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*{{cite journal | author=Nicolaides NC, Holroyd KJ, Ewart SL, ''et al.'' |title=Interleukin 9: a candidate gene for asthma. |journal=Proc. Natl. Acad. Sci. U.S.A. |volume=94 |issue= 24 |pages= 13175-80 |year= 1998 |pmid= 9371819 |doi= }} |
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*{{cite journal | author=Demoulin JB, Van Roost E, Stevens M, ''et al.'' |title=Distinct roles for STAT1, STAT3, and STAT5 in differentiation gene induction and apoptosis inhibition by interleukin-9. |journal=J. Biol. Chem. |volume=274 |issue= 36 |pages= 25855-61 |year= 1999 |pmid= 10464327 |doi= }} |
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*{{cite journal | author=Lejeune D, Demoulin JB, Renauld JC |title=Interleukin 9 induces expression of three cytokine signal inhibitors: cytokine-inducible SH2-containing protein, suppressor of cytokine signalling (SOCS)-2 and SOCS-3, but only SOCS-3 overexpression suppresses interleukin 9 signalling. |journal=Biochem. J. |volume=353 |issue= Pt 1 |pages= 109-116 |year= 2001 |pmid= 11115404 |doi= }} |
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*{{cite journal | author=Little FF, Cruikshank WW, Center DM |title=Il-9 stimulates release of chemotactic factors from human bronchial epithelial cells. |journal=Am. J. Respir. Cell Mol. Biol. |volume=25 |issue= 3 |pages= 347-52 |year= 2001 |pmid= 11588013 |doi= }} |
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*{{cite journal | author=Toda M, Tulic MK, Levitt RC, Hamid Q |title=A calcium-activated chloride channel (HCLCA1) is strongly related to IL-9 expression and mucus production in bronchial epithelium of patients with asthma. |journal=J. Allergy Clin. Immunol. |volume=109 |issue= 2 |pages= 246-50 |year= 2002 |pmid= 11842292 |doi= }} |
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*{{cite journal | author=Pilette C, Ouadrhiri Y, Van Snick J, ''et al.'' |title=IL-9 inhibits oxidative burst and TNF-alpha release in lipopolysaccharide-stimulated human monocytes through TGF-beta. |journal=J. Immunol. |volume=168 |issue= 8 |pages= 4103-11 |year= 2002 |pmid= 11937570 |doi= }} |
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}} |
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{{refend}} |
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== Discovery == |
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{{interleukins}} |
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IL-9 was first described in the late 1980s as a member of a growing number of [[cytokines]] that had [[Pleiotropy|pleiotropic]] functions in the [[immune]] system.IL-9 remains an understudied cytokine even though it has been allocated with many biological functions. It was first purified and characterized as a [[T cell]] and [[mast cell]] growth factor and termed as P40, based on their [[Molecular Weight|molecular weight]], or Mast cell growth-enhancing activity (MEA).The [[cloning]] and complete [[amino acid]] sequencing of P40 disclosed that it is structurally different from other [[T cell]]s growth factors. So, it was named IL-9 based on its biological effects on both [[Myeloid cells|myeloid]] and [[Lymphoid cells|lymphoid]] cells.<ref>{{cite journal | vauthors = Goswami R, Kaplan MH | title = A brief history of IL-9 | journal = Journal of Immunology | volume = 186 | issue = 6 | pages = 3283–8 | date = March 2011 | pmid = 21368237 | pmc = 3074408 | doi = 10.4049/jimmunol.1003049 }}</ref> |
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The identification and [[cloning]] was first done by Yang and colleagues as a [[mitogenic factor]] for a human megakaryoblastic leukemia. The same human [[cDNA]] was isolated again by cross-hybridization with the mouse IL-9 probe.<ref name="Renauld 1995 287–303">{{cite book|title=Cytokines: Interleukins and Their Receptors|volume = 80|last=Renauld|first=Jean-Christophe |date=1995|publisher=Springer, Boston, MA|isbn=9781461285281|series=Cancer Treatment and Research|pages=287–303|language=en|doi=10.1007/978-1-4613-1241-3_11|chapter = Interleukin-9: Structural characteristics and biologic properties|pmid = 8821582}}</ref> |
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== Gene location == |
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The human IL-9 gene is located on the long arm of human [[Chromosome 5 (human)|chromosome 5]] at band 5q31-32, a region which is not found in a number of patients with acquired [[chromosome 5q deletion syndrome]].<ref name="pmid1901233">{{cite journal | vauthors = Kelleher K, Bean K, Clark SC, Leung WY, Yang-Feng TL, Chen JW, Lin PF, Luo W, Yang YC | title = Human interleukin-9: genomic sequence, chromosomal location, and sequences essential for its expression in human T-cell leukemia virus (HTLV)-I-transformed human T cells | journal = Blood | volume = 77 | issue = 7 | pages = 1436–41 | year = 1991 | pmid = 1901233 | doi = 10.1182/blood.V77.7.1436.1436| url = http://www.bloodjournal.org/content/bloodjournal/77/7/1436.full.pdf | doi-access = free }}</ref> |
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== Protein structure == |
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Human IL-9 [[protein]] sequence contains 144 residues with a typical [[signal peptide]] of 18 [[amino acid]]s. There is also the presence of 9 [[cysteine]]s in mature [[polypeptide]] and 4 [[N-linked glycosylation]] sites.<ref name="Renauld 1995 287–303"/> Until recently, IL-9 was thought to be evolutionary related to IL-7.<ref>{{cite journal|last1=Boulay|first1=J. L.|last2=Paul|first2=W. E.|date=1993-09-01|title=Hematopoietin sub-family classification based on size, gene organization and sequence homology|journal=Current Biology|volume=3|issue=9|pages=573–581|issn=0960-9822|pmid=15335670|doi=10.1016/0960-9822(93)90002-6|bibcode=1993CBio....3..573B |s2cid=42479456}}</ref> However, we know now that IL-9 is closer to IL-2 and IL-15 than to IL-7,<ref>{{cite journal|last=Reche|first=Pedro A.|date=2019-02-01|title=The tertiary structure of γc cytokines dictates receptor sharing|journal=Cytokine|volume=116|pages=161–168|doi=10.1016/j.cyto.2019.01.007|issn=1096-0023|pmid=30716660|s2cid=73449371}}</ref> at both the tertiary and amino acid sequence levels. |
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== Production == |
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[[Interleukin 33]] (IL-33) induces IL-9 expression and secretion in [[T cell]]s, which was confirmed by the results obtained in mice by using Human ''[[in vitro]]''<nowiki/> system.<ref>{{cite journal | vauthors = Humphreys NE, Xu D, Hepworth MR, Liew FY, Grencis RK | title = IL-33, a potent inducer of adaptive immunity to intestinal nematodes | journal = Journal of Immunology | volume = 180 | issue = 4 | pages = 2443–9 | date = February 2008 | pmid = 18250453 | doi=10.4049/jimmunol.180.4.2443| doi-access = free }}</ref> Whereas the report of others confirms that [[TGF-b|TGF-β]] is an essential factor for IL-9 induction.<ref>{{cite journal | vauthors = Beriou G, Bradshaw EM, Lozano E, Costantino CM, Hastings WD, Orban T, Elyaman W, Khoury SJ, Kuchroo VK, Baecher-Allan C, Hafler DA | title = TGF-beta induces IL-9 production from human Th17 cells | journal = Journal of Immunology | volume = 185 | issue = 1 | pages = 46–54 | date = July 2010 | pmid = 20498357 | pmc = 2936106 | doi = 10.4049/jimmunol.1000356 }}</ref> For the first time (Lars Blom, Britta C. Poulsen, Bettina M. Jensen, Anker Hansen and Lars K. Poulsen published a journal online in 2011 Jul 6),indicating that TGF-β may be important for production of IL-9 but it is not only the definite requirement for IL-9 induction, since [[Cell culture|cultures]] with IL-33 without TGF-β have noticeably increased secretion of IL-9, suggesting an important role of IL-33, even though that the effect was not found significant on the [[gene]] level.<ref>{{cite journal | vauthors = Blom L, Poulsen BC, Jensen BM, Hansen A, Poulsen LK | title = IL-33 induces IL-9 production in human CD4+ T cells and basophils | journal = PLOS ONE | volume = 6 | issue = 7 | pages = e21695 | date = 2011-07-06 | pmid = 21765905 | pmc = 3130774 | doi = 10.1371/journal.pone.0021695 | bibcode = 2011PLoSO...621695B | doi-access = free }}</ref> |
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[[File:IL9R_HUMAN.png|thumb|Interleukin-9 receptor]] |
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[[File:IL_9_functions.large.jpg|thumb|IL-9 has different direct and indirect effects on multiple cell types that affect the development of immunity and inflammation.|245x245px]] |
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== IL-9 expression == |
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The analysis of IL-9 expression in different types of [[tumours]] such as Large cell anaplastic lymphoma (LCAL) and [[Hodgkin's Disease|Hodgkin's Disease (HD)]] by [[Northern blot|Northern blot analysis]] and [[in situ hybridization]] has showed that IL-9 is not involved as an autocrine growth factor in the [[pathogenesis]] of most [[B-cell lymphomas|B and T-cell lymphomas]], but it may have a part in HD and LCAL autocrine growth. |
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The further investigation could be done to conclude another probability, that, the [[in vivo]] [[Gene expression|overexpression]] of IL-9 might show the unique [[symptom]]s related to [[eosinophilia]] which was recently reported for [[Interleukin 5]] positive cases of HD.<ref name="pmid1908723">{{cite journal | vauthors = Merz H, Houssiau FA, Orscheschek K, Renauld JC, Fliedner A, Herin M, Noel H, Kadin M, Mueller-Hermelink HK, Van Snick J | title = Interleukin-9 expression in human malignant lymphomas: unique association with Hodgkin's disease and large cell anaplastic lymphoma | journal = Blood | volume = 78 | issue = 5 | pages = 1311–7 | year = 1991 | pmid = 1908723 | doi = 10.1182/blood.V78.5.1311.1311| url = http://www.bloodjournal.org/content/78/5/1311 | doi-access = free }}</ref> |
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IL-9 was found to be the first physiological stimulus triggering ''[[BCL3]]'' expression in [[T cell]]s and [[mast cell]]s by the analysis done in mouse.<ref name="pmid10361130">{{cite journal | vauthors = Richard M, Louahed J, Demoulin JB, Renauld JC | title = Interleukin-9 regulates NF-kappaB activity through BCL3 gene induction | journal = Blood | volume = 93 | issue = 12 | pages = 4318–27 | year = 1999 | pmid = 10361130 | doi = 10.1182/blood.V93.12.4318| url = http://www.bloodjournal.org/content/93/12/4318 }}</ref> |
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== References == |
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{{reflist|33em}} |
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== Further reading == |
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{{Refbegin|33em}} |
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* {{cite book | vauthors = Renauld JC, Houssiau F, Louahed J, Vink A, Van Snick J, Uyttenhove C | title = Advances in Immunology Volume 54 | chapter = Interleukin-9 | volume = 54 | pages = 79–97 | year = 1993 | pmid = 8379467 | doi = 10.1016/S0065-2776(08)60533-7 | isbn = 978-0-12-022454-8 }} |
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* {{cite journal | vauthors = Knoops L, Renauld JC | title = IL-9 and its receptor: from signal transduction to tumorigenesis | journal = Growth Factors | volume = 22 | issue = 4 | pages = 207–15 | date = December 2004 | pmid = 15621723 | doi = 10.1080/08977190410001720879 | s2cid = 40523692 }} |
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* {{cite journal | vauthors = Modi WS, Pollock DD, Mock BA, Banner C, Renauld JC, Van Snick J | title = Regional localization of the human glutaminase (GLS) and interleukin-9 (IL9) genes by in situ hybridization | journal = Cytogenetics and Cell Genetics | volume = 57 | issue = 2–3 | pages = 114–6 | year = 1991 | pmid = 1680606 | doi = 10.1159/000133126 }} |
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* {{cite journal | vauthors = Kelleher K, Bean K, Clark SC, Leung WY, Yang-Feng TL, Chen JW, Lin PF, Luo W, Yang YC | title = Human interleukin-9: genomic sequence, chromosomal location, and sequences essential for its expression in human T-cell leukemia virus (HTLV)-I-transformed human T cells | journal = Blood | volume = 77 | issue = 7 | pages = 1436–41 | date = April 1991 | pmid = 1901233 | doi = 10.1182/blood.V77.7.1436.1436| doi-access = free }} |
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* {{cite journal | vauthors = Holbrook ST, Ohls RK, Schibler KR, Yang YC, Christensen RD | title = Effect of interleukin-9 on clonogenic maturation and cell-cycle status of fetal and adult hematopoietic progenitors | journal = Blood | volume = 77 | issue = 10 | pages = 2129–34 | date = May 1991 | pmid = 1903074 | doi = 10.1182/blood.V77.10.2129.2129| doi-access = free }} |
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* {{cite journal | vauthors = Merz H, Houssiau FA, Orscheschek K, Renauld JC, Fliedner A, Herin M, Noel H, Kadin M, Mueller-Hermelink HK, Van Snick J | title = Interleukin-9 expression in human malignant lymphomas: unique association with Hodgkin's disease and large cell anaplastic lymphoma | journal = Blood | volume = 78 | issue = 5 | pages = 1311–7 | date = September 1991 | pmid = 1908723 | doi = 10.1182/blood.V78.5.1311.1311| doi-access = free }} |
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* {{cite journal | vauthors = Renauld JC, Goethals A, Houssiau F, Merz H, Van Roost E, Van Snick J | title = Human P40/IL-9. Expression in activated CD4+ T cells, genomic organization, and comparison with the mouse gene | journal = Journal of Immunology | volume = 144 | issue = 11 | pages = 4235–41 | date = June 1990 | doi = 10.4049/jimmunol.144.11.4235 | pmid = 1971295 | s2cid = 30151082 }} |
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* {{cite journal | vauthors = Renauld JC, Goethals A, Houssiau F, Van Roost E, Van Snick J | title = Cloning and expression of a cDNA for the human homolog of mouse T cell and mast cell growth factor P40 | journal = Cytokine | volume = 2 | issue = 1 | pages = 9–12 | date = January 1990 | pmid = 2129501 | doi = 10.1016/1043-4666(90)90037-T | hdl = 2078.1/11464 }} |
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* {{cite journal | vauthors = Yang YC, Ricciardi S, Ciarletta A, Calvetti J, Kelleher K, Clark SC | title = Expression cloning of cDNA encoding a novel human hematopoietic growth factor: human homologue of murine T-cell growth factor P40 | journal = Blood | volume = 74 | issue = 6 | pages = 1880–4 | date = November 1989 | pmid = 2508790 | doi = 10.1182/blood.V74.6.1880.1880| doi-access = free }} |
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* {{cite journal | vauthors = Yin T, Keller SR, Quelle FW, Witthuhn BA, Tsang ML, Lienhard GE, Ihle JN, Yang YC | title = Interleukin-9 induces tyrosine phosphorylation of insulin receptor substrate-1 via JAK tyrosine kinases | journal = The Journal of Biological Chemistry | volume = 270 | issue = 35 | pages = 20497–502 | date = September 1995 | pmid = 7544789 | doi = 10.1074/jbc.270.35.20497 | doi-access = free }} |
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* {{cite journal | vauthors = Postma DS, Bleecker ER, Amelung PJ, Holroyd KJ, Xu J, Panhuysen CI, Meyers DA, Levitt RC | title = Genetic susceptibility to asthma--bronchial hyperresponsiveness coinherited with a major gene for atopy | journal = The New England Journal of Medicine | volume = 333 | issue = 14 | pages = 894–900 | date = October 1995 | pmid = 7666875 | doi = 10.1056/NEJM199510053331402 | doi-access =free }} |
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* {{cite journal | vauthors = Le Beau MM, Espinosa R, Neuman WL, Stock W, Roulston D, Larson RA, Keinanen M, Westbrook CA | title = Cytogenetic and molecular delineation of the smallest commonly deleted region of chromosome 5 in malignant myeloid diseases | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 90 | issue = 12 | pages = 5484–8 | date = June 1993 | pmid = 8516290 | pmc = 46745 | doi = 10.1073/pnas.90.12.5484 | bibcode = 1993PNAS...90.5484B | doi-access = free }} |
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* {{cite journal | vauthors = Demoulin JB, Uyttenhove C, Van Roost E, DeLestré B, Donckers D, Van Snick J, Renauld JC | title = A single tyrosine of the interleukin-9 (IL-9) receptor is required for STAT activation, antiapoptotic activity, and growth regulation by IL-9 | journal = Molecular and Cellular Biology | volume = 16 | issue = 9 | pages = 4710–6 | date = September 1996 | pmid = 8756628 | pmc = 231471 | doi = 10.1128/mcb.16.9.4710}} |
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* {{cite journal | vauthors = Nicolaides NC, Holroyd KJ, Ewart SL, Eleff SM, Kiser MB, Dragwa CR, Sullivan CD, Grasso L, Zhang LY, Messler CJ, Zhou T, Kleeberger SR, Buetow KH, Levitt RC | title = Interleukin 9: a candidate gene for asthma | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 94 | issue = 24 | pages = 13175–80 | date = November 1997 | pmid = 9371819 | pmc = 24282 | doi = 10.1073/pnas.94.24.13175 | bibcode = 1997PNAS...9413175N | doi-access = free }} |
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* {{cite journal | vauthors = Demoulin JB, Van Roost E, Stevens M, Groner B, Renauld JC | title = Distinct roles for STAT1, STAT3, and STAT5 in differentiation gene induction and apoptosis inhibition by interleukin-9 | journal = The Journal of Biological Chemistry | volume = 274 | issue = 36 | pages = 25855–61 | date = September 1999 | pmid = 10464327 | doi = 10.1074/jbc.274.36.25855 | doi-access = free}} |
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* {{cite journal | vauthors = Lejeune D, Demoulin JB, Renauld JC | title = Interleukin 9 induces expression of three cytokine signal inhibitors: cytokine-inducible SH2-containing protein, suppressor of cytokine signalling (SOCS)-2 and SOCS-3, but only SOCS-3 overexpression suppresses interleukin 9 signalling | journal = The Biochemical Journal | volume = 353 | issue = Pt 1 | pages = 109–116 | date = January 2001 | pmid = 11115404 | pmc = 1221548 | doi = 10.1042/0264-6021:3530109 }} |
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* {{cite journal | vauthors = Little FF, Cruikshank WW, Center DM | title = Il-9 stimulates release of chemotactic factors from human bronchial epithelial cells | journal = American Journal of Respiratory Cell and Molecular Biology | volume = 25 | issue = 3 | pages = 347–52 | date = September 2001 | pmid = 11588013 | doi = 10.1165/ajrcmb.25.3.4349 }} |
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* {{cite journal | vauthors = Toda M, Tulic MK, Levitt RC, Hamid Q | title = A calcium-activated chloride channel (HCLCA1) is strongly related to IL-9 expression and mucus production in bronchial epithelium of patients with asthma | journal = The Journal of Allergy and Clinical Immunology | volume = 109 | issue = 2 | pages = 246–50 | date = February 2002 | pmid = 11842292 | doi = 10.1067/mai.2002.121555 | doi-access = free }} |
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* {{cite journal | vauthors = Pilette C, Ouadrhiri Y, Van Snick J, Renauld JC, Staquet P, Vaerman JP, Sibille Y | title = IL-9 inhibits oxidative burst and TNF-alpha release in lipopolysaccharide-stimulated human monocytes through TGF-beta | journal = Journal of Immunology | volume = 168 | issue = 8 | pages = 4103–11 | date = April 2002 | pmid = 11937570 | doi = 10.4049/jimmunol.168.8.4103 | doi-access = free }} |
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{{refend}} |
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{{gene-5-stub}} |
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{{Interleukins}} |
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{{Interleukin receptor modulators}} |
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{{immunology-stub}} |
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[[Category:Interleukins]] |
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[[pl:Interleukina 9]] |
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