Ver o código fonte de "Factor de crecemento de fibroblastos ácido"

{{Xene-proteína
|nome           = FGF1
|nomes          = FGF1, AFGF, ECGF, ECGF-beta, ECGFA, ECGFB, FGF-1, FGF-alfa, FGFA, GLIO703, HBGF-1, HBGF1, factor de crecemento de fibroblastos 1
|imaxe           = Protein FGF1 PDB 1afc.png
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|Cromosoma        = 5
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|Banda            = 31.3
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|Homologene       = 625
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|OMIM             = 131220
|GeneAtlas_image1 =PBB GE FGF1 208240 s at fs.png
|GeneAtlas_image2 =PBB GE FGF1 205117 at fs.png
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|NCBI             =
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<!-- Ortoloxía -->
|Hs_EntrezGene =2246
|Hs_Ensembl =ENSG00000113578
|Hs_RefseqmRNA =NM_000800
|Hs_RefseqProtein =NP_000791
|Hs_GenLoc_db =hg5
|Hs_GenLoc_chr = 5
|Hs_GenLoc_start =142592178
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}}
O '''factor de crecemento de fibroblastos ácido''' ('''aFGF'''), tamén chamado '''FGF1''' ou '''FGF-α''' é un [[factor de crecemento]] e [[proteína de sinalización]] codificado polo [[xene]] ''FGF1'' do [[cromosoma 5]] humano.<ref name="pmid1697263">{{Cita publicación periódica | vauthors = Dionne CA, Crumley G, Bellot F, Kaplow JM, Searfoss G, Ruta M, Burgess WH, Jaye M, Schlessinger J | title = Cloning and expression of two distinct high-affinity receptors cross-reacting with acidic and basic fibroblast growth factors | journal = The EMBO Journal | volume = 9 | issue = 9 | pages = 2685–92 | date = September 1990 | pmid = 1697263 | pmc = 551973 | doi =  }}</ref><ref name="pmid3523756">{{Cita publicación periódica | vauthors = Jaye M, Howk R, Burgess W, Ricca GA, Chiu IM, Ravera MW, O'Brien SJ, Modi WS, Maciag T, Drohan WN | title = Human endothelial cell growth factor: cloning, nucleotide sequence, and chromosome localization | journal = Science | volume = 233 | issue = 4763 | pages = 541–5 | date = August 1986 | pmid = 3523756 | pmc =  | doi = 10.1126/science.3523756 }}</ref> É sintetizado como un polipéptido de 155 [[aminoácido]]s, cuxa forma madura é unha proteína non [[glicosilación|glicosilada]] de 17-18 kDa. A proteína [[factor de crecemento de fibroblastos]] foi purificada primeiramente en 1975, pero pouco despois, usando diferentes condicións, illáronse o FGF ácido, o factor de crecemento de heparina 1 e o [[ECGF1|factor de crecemento endotelial-1]].<ref>{{Cita publicación periódica | vauthors = Burgess WH, Maciag T | title = The heparin-binding (fibroblast) growth factor family of proteins | url = https://archive.org/details/sim_annual-review-of-biochemistry_1989_58/page/575 | journal = Annual Review of Biochemistry | volume = 58 | pages = 575–606 | date = 1989 | pmid = 2549857 | doi = 10.1146/annurev.bi.58.070189.003043 }}</ref> A secuenciación dos xenes revelou que o FGF1 era un membro da [[factor de crecemento de fibroblastos|familia de FGF]] de proteínas.

O FGF1 non ten unha secuencia sinal clara, polo que non é segregada a través das vías clásicas, pero non parece formar un dímero ligado por [[ponte disulfuro]] dentro das células, que se asocia cun complexo de proteínas na membrana plasmática (incluíndo [[S100A13]] e [[Sinaptotagmina 1|Syt1]]) que despois axudan a pasala a través da membrana ata o exterior da célula.<ref>{{Cita publicación periódica | vauthors = Tarantini F, Gamble S, Jackson A, Maciag T | title = The cysteine residue responsible for the release of fibroblast growth factor-1 residues in a domain independent of the domain for phosphatidylserine binding | journal = The Journal of Biological Chemistry | volume = 270 | issue = 49 | pages = 29039–42 | date = December 1995 | pmid = 7493920 | doi = 10.1074/jbc.270.49.29039 | url = http://www.jbc.org/content/270/49/29039.long }}</ref><ref name="Prudovsky&Maciag">{{Cita publicación periódica | vauthors = Prudovsky I, Bagala C, Tarantini F, Mandinova A, Soldi R, Bellum S, Maciag T | title = The intracellular translocation of the components of the fibroblast growth factor 1 release complex precedes their assembly prior to export | journal = The Journal of Cell Biology | volume = 158 | issue = 2 | pages = 201–8 | date = July 2002 | pmid = 12135982 | pmc = 2173119 | doi = 10.1083/jcb.200203084 | url = http://jcb.rupress.org/content/158/2/201.long }}</ref> Unha vez que está nas condicións [[redox|redutoras]] dos tecidos circundantes, o dímero disóciase formando o FGF1 [[monómero]] que pode entrar na circulación sistémica ou ser secuestrado nos tecidos uníndose a [[proteoglicano de heparán sulfato|proteoglicanos de heparán sulfato]] da [[matriz extracelular]]. O FGF1 pode despois unirse e exercer os seus efectos por medio de proteínas específicas [[receptor do factor de crecemento de fibroblastos|receptoras do factor de crecemento de fibroblastos (FGFR)]], as cales constitúen unha familia de moléculas estreitamente relacionadas.<ref name="pmid 24780002">{{Cita publicación periódica | vauthors = Coleman SJ, Bruce C, Chioni AM, Kocher HM, Grose RP | title = The ins and outs of fibroblast growth factor receptor signalling | journal = Clinical Science | volume = 127 | issue = 4 | pages = 217–31 | date = August 2014 | pmid = 24780002 | doi = 10.1042/CS20140100 | url = http://www.clinsci.org/content/127/4/217 }}</ref>

Ademais da súa actividade extracelular, o FGF1 pode tamén funcionar intracelularmente. A proteína ten unha [[secuencia de localización nuclear]] (NLS), pero a ruta que segue o FGF1 para chegar ao [[núcleo celular|núcleo]] non está clara e parece que cómpre que se una a algún tipo de [[receptor da superficie celular]], seguido da súa internalización e translocación ao núcleo, onde pode interaccionar con [[isoforma]]s nucleares de FGFR.<ref name="pmid 24780002" /> Isto é diferente do que ocorre co [[FGF2]], que tamén pode activar os FGFR nucleares, pero que ten variantes de [[splicing|empalme]] da proteína que nunca abandonan a célula e van directamente ao núcleo.

== Función ==

Os membros da familia do FGF posúen amplas actividades [[mitoxénica]]s e de supervivencia celular e están implicados en diversos procesos biolóxicos, como o [[desenvolvemento embrionario]], [[crecemento celular]], [[morfoxénese]], [[reparación de tecidos]] e o crecemento e invasión tumorais. Esta proteína funciona como un modificador da migración e proliferación de células endoteliais, así como un factor [[anxioxénese|anxioxénico]]. Actúa como [[mitóxeno]] para diversas células derivadas do mesoderma e neuroectoderma in vitro, así se pensa que está implicado na [[organoxénese]]. Describíronse tres variantes de [[empalme alternativo]] que codifican diferentes [[isoforma]]s.<ref>{{Cita web | title = Entrez Gene: FGF1 fibroblast growth factor 1 (acidic)| url = https://www.ncbi.nlm.nih.gov/sites/entrez?Db=gene&Cmd=ShowDetailView&TermToSearch=2246| access-date = }}</ref>

O FGF1 é multifuncional e ten moitos efectos. Como exemplo, en ratos con [[diabetes]] inducida pola dieta, que é un equivalente experimental da diabetes de tipo 2 humana, unha soa inxección da proteína FGF1 é dabondo para restablecer os niveis de azucre do sangue ata uns valores saudables durante máis de dous días.<ref>{{Cita publicación periódica | vauthors = Suh JM, Jonker JW, Ahmadian M, Goetz R, Lackey D, Osborn O, Huang Z, Liu W, Yoshihara E, van Dijk TH, Havinga R, Fan W, Yin YQ, Yu RT, Liddle C, Atkins AR, Olefsky JM, Mohammadi M, Downes M, Evans RM | title = Endocrinization of FGF1 produces a neomorphic and potent insulin sensitizer | journal = Nature | volume = 513 | issue = 7518 | pages = 436–9 | date = September 2014 | pmid = 25043058 | pmc = 4184286 | doi = 10.1038/nature13540 | laysummary = http://www.salk.edu/news/pressrelease_details.php?press_id=2037 | laysource = Salk Institute }}</ref>

== Interaccións ==

O FGF1 presenta [[interaccións proteína-proteína|interaccións]] con:
* [[CSNK2A2]]<ref name = pmid12145206/> 
* [[CSNK2B]]<ref name = pmid12145206>{{Cita publicación periódica | vauthors = Skjerpen CS, Nilsen T, Wesche J, Olsnes S | title = Binding of FGF-1 variants to protein kinase CK2 correlates with mitogenicity | journal = The EMBO Journal | volume = 21 | issue = 15 | pages = 4058–69 | date = August 2002 | pmid = 12145206 | pmc = 126148 | doi = 10.1093/emboj/cdf402 }}</ref> 
* [[Caseína quinase 2, alfa 1|CSNK2A1]]<ref name = pmid12145206/> 
* [[FIBP]]<ref name = pmid9806903>{{Cita publicación periódica | vauthors = Kolpakova E, Wiedłocha A, Stenmark H, Klingenberg O, Falnes PO, Olsnes S | title = Cloning of an intracellular protein that binds selectively to mitogenic acidic fibroblast growth factor | journal = The Biochemical Journal | volume = 336 ( Pt 1) | issue = 1 | pages = 213–22 | date = November 1998 | pmid = 9806903 | pmc = 1219860 | doi = 10.1042/bj3360213 }}</ref> 
* [[Receptor do factor de crecemento de fibroblastos 1|FGFR1]]<ref name = pmid11030354>{{Cita publicación periódica | vauthors = Schlessinger J, Plotnikov AN, Ibrahimi OA, Eliseenkova AV, Yeh BK, Yayon A, Linhardt RJ, Mohammadi M | title = Crystal structure of a ternary FGF-FGFR-heparin complex reveals a dual role for heparin in FGFR binding and dimerization | journal = Molecular Cell | volume = 6 | issue = 3 | pages = 743–50 | date = September 2000 | pmid = 11030354 | doi = 10.1016/s1097-2765(00)00073-3 }}</ref><ref name = pmid8576175>{{Cita publicación periódica | vauthors = Santos-Ocampo S, Colvin JS, Chellaiah A, Ornitz DM | title = Expression and biological activity of mouse fibroblast growth factor-9 | journal = The Journal of Biological Chemistry | volume = 271 | issue = 3 | pages = 1726–31 | date = January 1996 | pmid = 8576175 | doi = 10.1074/jbc.271.3.1726 }}</ref> 
* [[Receptor do factor de crecemento de fibroblastos 2|FGFR2]]<ref name = pmid8576175/><ref name = pmid10618369>{{Cita publicación periódica | vauthors = Stauber DJ, DiGabriele AD, Hendrickson WA | title = Structural interactions of fibroblast growth factor receptor with its ligands | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 97 | issue = 1 | pages = 49–54 | date = January 2000 | pmid = 10618369 | pmc = 26614 | doi = 10.1073/pnas.97.1.49 }}</ref><ref name = pmid11069186>{{Cita publicación periódica | vauthors = Pellegrini L, Burke DF, von Delft F, Mulloy B, Blundell TL | title = Crystal structure of fibroblast growth factor receptor ectodomain bound to ligand and heparin | journal = Nature | volume = 407 | issue = 6807 | pages = 1029–34 | date = October 2000 | pmid = 11069186 | doi = 10.1038/35039551 }}</ref> 
* [[Receptor do factor de crecemento de fibrblastos 3|FGFR3]]<ref name = pmid8576175/><ref name = pmid10574949>{{Cita publicación periódica | vauthors = Chellaiah A, Yuan W, Chellaiah M, Ornitz DM | title = Mapping ligand binding domains in chimeric fibroblast growth factor receptor molecules. Multiple regions determine ligand binding specificity | journal = The Journal of Biological Chemistry | volume = 274 | issue = 49 | pages = 34785–94 | date = December 1999 | pmid = 10574949 | doi = 10.1074/jbc.274.49.34785 }}</ref>
* [[Receptor do factor de c recemento de fibroblastos 4|FGFR4]]<ref name = pmid10736564>{{Cita publicación periódica | vauthors = Loo BB, Darwish KK, Vainikka SS, Saarikettu JJ, Vihko PP, Hermonen JJ, Goldman AA, Alitalo KK, Jalkanen MM | title = Production and characterization of the extracellular domain of recombinant human fibroblast growth factor receptor 4 | url = https://archive.org/details/sim_international-journal-of-biochemistry-cell-biology_2000-05_32_5/page/489 | journal = The International Journal of Biochemistry & Cell Biology | volume = 32 | issue = 5 | pages = 489–97 | date = May 2000 | pmid = 10736564 | doi = 10.1016/S1357-2725(99)00145-4 }}</ref><ref name = pmid10336501>{{Cita publicación periódica | vauthors = Kan M, Wu X, Wang F, McKeehan WL | title = Specificity for fibroblast growth factors determined by heparan sulfate in a binary complex with the receptor kinase | journal = The Journal of Biological Chemistry | volume = 274 | issue = 22 | pages = 15947–52 | date = May 1999 | pmid = 10336501 | doi = 10.1074/jbc.274.22.15947 }}</ref> 
* [[HSPA9]]<ref name = pmid10510314>{{Cita publicación periódica | vauthors = Mizukoshi E, Suzuki M, Loupatov A, Uruno T, Hayashi H, Misono T, Kaul SC, Wadhwa R, Imamura T | title = Fibroblast growth factor-1 interacts with the glucose-regulated protein GRP75/mortalin | journal = The Biochemical Journal | volume = 343 Pt 2 | issue = 2 | pages = 461–6 | date = October 1999 | pmid = 10510314 | pmc = 1220575 | doi = 10.1042/0264-6021:3430461 }}</ref> 
* [[S100A13]]<ref name="Prudovsky&Maciag" /><ref name = "pmid9712836">{{Cita publicación periódica | vauthors = Mouta Carreira C, LaVallee TM, Tarantini F, Jackson A, Lathrop JT, Hampton B, Burgess WH, Maciag T | title = S100A13 is involved in the regulation of fibroblast growth factor-1 and p40 synaptotagmin-1 release in vitro | journal = The Journal of Biological Chemistry | volume = 273 | issue = 35 | pages = 22224–31 | date = August 1998 | pmid = 9712836 | doi = 10.1074/jbc.273.35.22224 }}</ref><ref name = pmid11432880>{{Cita publicación periódica | vauthors = Landriscina M, Bagalá C, Mandinova A, Soldi R, Micucci I, Bellum S, Prudovsky I, Maciag T | title = Copper induces the assembly of a multiprotein aggregate implicated in the release of fibroblast growth factor 1 in response to stress | journal = The Journal of Biological Chemistry | volume = 276 | issue = 27 | pages = 25549–57 | date = July 2001 | pmid = 11432880 | doi = 10.1074/jbc.M102925200 }}</ref> e
* [[Sinaptotagmina 1|Sinaptotagmina 1 (SYT1)]] <ref name="Prudovsky&Maciag" /><ref name=pmid9712836 />

== Notas ==
{{listaref}}

== Véxase tamén ==
=== Outros artigos ===
* [[Factor de crecemento de fibroblastos]]
* [[Factor de crecemento de fibroblastos básico]]

=== Bibliografía ===
* {{Cita publicación periódica | vauthors = Yu YL, Kha H, Golden JA, Migchielsen AA, Goetzl EJ, Turck CW | title = An acidic fibroblast growth factor protein generated by alternate splicing acts like an antagonist | journal = The Journal of Experimental Medicine | volume = 175 | issue = 4 | pages = 1073–80 | date = April 1992 | pmid = 1372643 | pmc = 2119192 | doi = 10.1084/jem.175.4.1073 }}
* {{Cita publicación periódica | vauthors = Chiu IM, Wang WP, Lehtoma K | title = Alternative splicing generates two forms of mRNA coding for human heparin-binding growth factor 1 | url = https://archive.org/details/sim_oncogene_1990-05_5_5/page/755 | journal = Oncogene | volume = 5 | issue = 5 | pages = 755–62 | date = May 1990 | pmid = 1693186 | doi =  }}
* {{Cita publicación periódica | vauthors = Zhu X, Komiya H, Chirino A, Faham S, Fox GM, Arakawa T, Hsu BT, Rees DC | title = Three-dimensional structures of acidic and basic fibroblast growth factors | journal = Science | volume = 251 | issue = 4989 | pages = 90–3 | date = January 1991 | pmid = 1702556 | doi = 10.1126/science.1702556 }}
* {{Cita publicación periódica | vauthors = Wang WP, Quick D, Balcerzak SP, Needleman SW, Chiu IM | title = Cloning and sequence analysis of the human acidic fibroblast growth factor gene and its preservation in leukemia patients | url = https://archive.org/details/sim_oncogene_1991-09_6_9/page/1521 | journal = Oncogene | volume = 6 | issue = 9 | pages = 1521–9 | date = September 1991 | pmid = 1717925 | doi =  }}
* {{Cita publicación periódica | vauthors = Wu DQ, Kan MK, Sato GH, Okamoto T, Sato JD | title = Characterization and molecular cloning of a putative binding protein for heparin-binding growth factors | journal = The Journal of Biological Chemistry | volume = 266 | issue = 25 | pages = 16778–85 | date = September 1991 | pmid = 1885605 | doi =  }}
* {{Cita publicación periódica | vauthors = Crumley G, Dionne CA, Jaye M | title = The gene for human acidic fibroblast growth factor encodes two upstream exons alternatively spliced to the first coding exon | journal = Biochemical and Biophysical Research Communications | volume = 171 | issue = 1 | pages = 7–13 | date = August 1990 | pmid = 2393407 | doi = 10.1016/0006-291X(90)91348-V }}
* {{Cita publicación periódica | vauthors = Harper JW, Strydom DJ, Lobb RR | title = Human class 1 heparin-binding growth factor: structure and homology to bovine acidic brain fibroblast growth factor | journal = Biochemistry | volume = 25 | issue = 14 | pages = 4097–103 | date = July 1986 | pmid = 2427112 | doi = 10.1021/bi00362a017 }}
* {{Cita publicación periódica | vauthors = Winkles JA, Friesel R, Burgess WH, Howk R, Mehlman T, Weinstein R, Maciag T | title = Human vascular smooth muscle cells both express and respond to heparin-binding growth factor I (endothelial cell growth factor) | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 84 | issue = 20 | pages = 7124–8 | date = October 1987 | pmid = 2444975 | pmc = 299242 | doi = 10.1073/pnas.84.20.7124 }}
* {{Cita publicación periódica | vauthors = Wang WP, Lehtoma K, Varban ML, Krishnan I, Chiu IM | title = Cloning of the gene coding for human class 1 heparin-binding growth factor and its expression in fetal tissues | journal = Molecular and Cellular Biology | volume = 9 | issue = 6 | pages = 2387–95 | date = June 1989 | pmid = 2474753 | pmc = 362312 | doi =  }}
* {{Cita publicación periódica | vauthors = Mergia A, Tischer E, Graves D, Tumolo A, Miller J, Gospodarowicz D, Abraham JA, Shipley GD, Fiddes JC | title = Structural analysis of the gene for human acidic fibroblast growth factor | journal = Biochemical and Biophysical Research Communications | volume = 164 | issue = 3 | pages = 1121–9 | date = November 1989 | pmid = 2590193 | doi = 10.1016/0006-291X(89)91785-3 }}
* {{Cita publicación periódica | vauthors = Gimenez-Gallego G, Conn G, Hatcher VB, Thomas KA | title = The complete amino acid sequence of human brain-derived acidic fibroblast growth factor | journal = Biochemical and Biophysical Research Communications | volume = 138 | issue = 2 | pages = 611–7 | date = July 1986 | pmid = 3527167 | doi = 10.1016/S0006-291X(86)80540-X }}
* {{Cita publicación periódica | vauthors = Gautschi P, Fràter-Schröder M, Böhlen P | title = Partial molecular characterization of endothelial cell mitogens from human brain: acidic and basic fibroblast growth factors | journal = FEBS Letters | volume = 204 | issue = 2 | pages = 203–7 | date = August 1986 | pmid = 3732516 | doi = 10.1016/0014-5793(86)80812-2 }}
* {{Cita publicación periódica | vauthors = Gautschi-Sova P, Müller T, Böhlen P | title = Amino acid sequence of human acidic fibroblast growth factor | journal = Biochemical and Biophysical Research Communications | volume = 140 | issue = 3 | pages = 874–80 | date = November 1986 | pmid = 3778488 | doi = 10.1016/0006-291X(86)90716-3 }}
* {{Cita publicación periódica | vauthors = Gimenez-Gallego G, Conn G, Hatcher VB, Thomas KA | title = Human brain-derived acidic and basic fibroblast growth factors: amino terminal sequences and specific mitogenic activities | journal = Biochemical and Biophysical Research Communications | volume = 135 | issue = 2 | pages = 541–8 | date = March 1986 | pmid = 3964259 | doi = 10.1016/0006-291X(86)90028-8 }}
* {{Cita publicación periódica | vauthors = Zhao XM, Yeoh TK, Hiebert M, Frist WH, Miller GG | title = The expression of acidic fibroblast growth factor (heparin-binding growth factor-1) and cytokine genes in human cardiac allografts and T cells | journal = Transplantation | volume = 56 | issue = 5 | pages = 1177–82 | date = November 1993 | pmid = 7504343 | doi = 10.1097/00007890-199311000-00025 }}
* {{Cita publicación periódica | vauthors = Pineda-Lucena A, Jiménez MA, Nieto JL, Santoro J, Rico M, Giménez-Gallego G | title = 1H-NMR assignment and solution structure of human acidic fibroblast growth factor activated by inositol hexasulfate | journal = Journal of Molecular Biology | volume = 242 | issue = 1 | pages = 81–98 | date = September 1994 | pmid = 7521397 | doi = 10.1006/jmbi.1994.1558 }}
* {{Cita publicación periódica | vauthors = Chotani MA, Payson RA, Winkles JA, Chiu IM | title = Human fibroblast growth factor 1 gene expression in vascular smooth muscle cells is modulated via an alternate promoter in response to serum and phorbol ester | journal = Nucleic Acids Research | volume = 23 | issue = 3 | pages = 434–41 | date = February 1995 | pmid = 7533902 | pmc = 306694 | doi = 10.1093/nar/23.3.434 }}
* {{Cita publicación periódica | vauthors = Opalenik SR, Shin JT, Wehby JN, Mahesh VK, Thompson JA | title = The HIV-1 TAT protein induces the expression and extracellular appearance of acidic fibroblast growth factor | journal = The Journal of Biological Chemistry | volume = 270 | issue = 29 | pages = 17457–67 | date = July 1995 | pmid = 7542239 | doi = 10.1074/jbc.270.29.17457 }}
* {{Cita publicación periódica | vauthors = Myers RL, Payson RA, Chotani MA, Deaven LL, Chiu IM | title = Gene structure and differential expression of acidic fibroblast growth factor mRNA: identification and distribution of four different transcripts | url = https://archive.org/details/sim_oncogene_1993-02_8_2/page/341 | journal = Oncogene | volume = 8 | issue = 2 | pages = 341–9 | date = February 1993 | pmid = 7678925 | doi =  }}

{{control de autoridades}}

[[Categoría:Factores de crecemento]]