title: Insights into the key roles of proteoglycans in breast cancer biology and translational medicine.
abstract: Proteoglycans control numerous normal and pathological processes, among which are morphogenesis, tissue repair, inflammation, vascularization and cancer metastasis. During tumor development and growth, proteoglycan expression is markedly modified in the tumor microenvironment. Altered expression of proteoglycans on tumor and stromal cell membranes affects cancer cell signaling, growth and survival, cell adhesion, migration and angiogenesis. Despite the high complexity and heterogeneity of breast cancer, the rapid evolution in our knowledge that proteoglycans are among the key players in the breast tumor microenvironment suggests their potential as pharmacological targets in this type of cancer. It has been recently suggested that pharmacological treatment may target proteoglycan metabolism, their utilization as targets for immunotherapy or their direct use as therapeutic agents. The diversity inherent in the proteoglycans that will be presented herein provides the potential for multiple layers of regulation of breast tumor behavior. This review summarizes recent developments concerning the biology of selected proteoglycans in breast cancer, and presents potential targeted therapeutic approaches based on their novel key roles in breast cancer.
authors.affiliation: Biochemistry, Biochemical Analysis & Matrix Pathobiology Research Group, Laboratory of Biochemistry, Department of Chemistry, University of Patras, 26500 Patras, Greece.
authors.forename: Achilleas D
authors.initials: AD
authors.lastname: Theocharis
authors.affiliation: Biochemistry, Biochemical Analysis & Matrix Pathobiology Research Group, Laboratory of Biochemistry, Department of Chemistry, University of Patras, 26500 Patras, Greece.
authors.forename: Spyros S
authors.initials: SS
authors.lastname: Skandalis
authors.affiliation: Department of Pathology, Anatomy and Cell Biology and the Cancer Cell Biology and Signaling Program, Kimmel Cancer Center, Thomas Jefferson University, Philadelphia, PA 19107, USA.
authors.forename: Thomas
authors.initials: T
authors.lastname: Neill
authors.affiliation: Department of Biomedical Sciences and Biotech Research & Innovation Center, University of Copenhagen, Denmark.
authors.forename: Hinke A B
authors.initials: HA
authors.lastname: Multhaupt
authors.affiliation: University of Frankfurt, Institute of Pharmacology and Toxicology, Theodor-Stern Kai 7, Frankfurt 60590, Germany.
authors.forename: Mario
authors.initials: M
authors.lastname: Hubo
authors.affiliation: University of Frankfurt, Institute of Pharmacology and Toxicology, Theodor-Stern Kai 7, Frankfurt 60590, Germany.
authors.forename: Helena
authors.initials: H
authors.lastname: Frey
authors.affiliation: Department of Biomedical Sciences and Biotech Research & Innovation Center, University of Copenhagen, Denmark.
authors.forename: Sandeep
authors.initials: S
authors.lastname: Gopal
authors.affiliation: Department of Biomedical Sciences and Biotech Research & Innovation Center, University of Copenhagen, Denmark.
authors.forename: Angélica
authors.initials: A
authors.lastname: Gomes
authors.affiliation: Department of Biomedical Sciences and Biotech Research & Innovation Center, University of Copenhagen, Denmark.
authors.forename: Nikos
authors.initials: N
authors.lastname: Afratis
authors.affiliation: Department of Biomedical Sciences and Biotech Research & Innovation Center, University of Copenhagen, Denmark.
authors.forename: Hooi Ching
authors.initials: HC
authors.lastname: Lim
authors.affiliation: Department of Biomedical Sciences and Biotech Research & Innovation Center, University of Copenhagen, Denmark.
authors.forename: John R
authors.initials: JR
authors.lastname: Couchman
authors.affiliation: Department of Biological Sciences, Sunnybrook Research Institute and Department of Medical Biophysics, University of Toronto, Canada.
authors.forename: Jorge
authors.initials: J
authors.lastname: Filmus
authors.affiliation: University of Alabama at Birmingham, Department of Pathology, UAB Comprehensive Cancer Center, 1720 2nd Ave. S, WTI 602B, Birmingham, AL 35294, USA.
authors.forename: Ralph D
authors.initials: RD
authors.lastname: Sanderson
authors.affiliation: University of Frankfurt, Institute of Pharmacology and Toxicology, Theodor-Stern Kai 7, Frankfurt 60590, Germany.
authors.forename: Liliana
authors.initials: L
authors.lastname: Schaefer
authors.affiliation: Department of Pathology, Anatomy and Cell Biology and the Cancer Cell Biology and Signaling Program, Kimmel Cancer Center, Thomas Jefferson University, Philadelphia, PA 19107, USA.
authors.forename: Renato V
authors.initials: RV
authors.lastname: Iozzo
authors.affiliation: Biochemistry, Biochemical Analysis & Matrix Pathobiology Research Group, Laboratory of Biochemistry, Department of Chemistry, University of Patras, 26500 Patras, Greece. Electronic address: n.k.karamanos@upatras.gr.
authors.forename: Nikos K
authors.initials: NK
authors.lastname: Karamanos
mesh_terms: D001943:Breast Neoplasms / Q000235:genetics* / Q000473:pathology; D005260:Female; D015972:Gene Expression Regulation, Neoplastic; D006801:Humans; D058990:Molecular Targeted Therapy; D009389:Neovascularization, Pathologic / Q000188:drug therapy / Q000235:genetics* / Q000473:pathology; D011509:Proteoglycans / Q000037:antagonists & inhibitors / Q000096:biosynthesis* / Q000627:therapeutic use; D015398:Signal Transduction / Q000235:genetics; D057170:Translational Research, Biomedical; D059016:Tumor Microenvironment / Q000235:genetics
chemical_list: D011509:Proteoglycans
keywords: Breast cancer; Decorin; Glypicans; Proteoglycans; Serglycin; Syndecans
references.citation: Cancer Res. 2012 Mar 15;72(6):1384-94
references.pmid: 22282653
references.citation: J Biol Chem. 2014 Dec 5;289(49):34089-103
references.pmid: 25320080
references.citation: Neoplasia. 2009 Oct;11(10):1042-53
references.pmid: 19794963
references.citation: Hepatology. 2014 Aug;60(2):576-87
references.pmid: 24492943
references.citation: Dev Cell. 2013 Mar 11;24(5):472-85
references.pmid: 23453597
references.citation: Gynecol Oncol. 2005 Oct;99(1):126-34
references.pmid: 16023706
references.citation: J Clin Invest. 2001 Apr;107(7):845-52
references.pmid: 11285303
references.citation: J Clin Invest. 2001 Jan;107(2):R9-R14
references.pmid: 11160142
references.citation: Cancer Res. 2011 Mar 1;71(5):1550-60
references.pmid: 21343399
references.citation: Cancer Res. 2004 Jan 15;64(2):612-21
references.pmid: 14744776
references.citation: Biochimie. 2006 Sep;88(9):1135-43
references.pmid: 16690200
references.citation: J Cell Biol. 2005 Nov 21;171(4):729-38
references.pmid: 16286510
references.citation: J Cell Sci. 2003 Dec 1;116(Pt 23):4857-64
references.pmid: 14600270
references.citation: Biochim Biophys Acta. 2012 Jul;1820(7):1009-19
references.pmid: 22542782
references.citation: FEBS J. 2013 May;280(10):2165-79
references.pmid: 23350913
references.citation: Eur J Cancer. 2004 Jun;40(9):1373-82
references.pmid: 15177497
references.citation: Nat Genet. 2000 Jul;25(3):329-32
references.pmid: 10888884
references.citation: Front Oncol. 2014 Feb 03;4:4
references.pmid: 24551591
references.citation: Cell. 2012 Dec 21;151(7):1542-56
references.pmid: 23260141
references.citation: Cancer Res. 2008 Nov 15;68(22):9558-65
references.pmid: 19010933
references.citation: Cell Tissue Res. 2010 Jan;339(1):237-46
references.pmid: 19513755
references.citation: Nat Med. 2010 Nov;16(11):1313-20
references.pmid: 21037586
references.citation: J Clin Invest. 1996 Feb 15;97(4):988-95
references.pmid: 8613553
references.citation: Cancer Cell. 2014 Apr 14;25(4):501-15
references.pmid: 24735924
references.citation: Anticancer Res. 2014 Jun;34(6):2797-800
references.pmid: 24922641
references.citation: Int J Cancer. 2006 Apr 1;118(7):1609-17
references.pmid: 16217746
references.citation: J Biol Chem. 1998 May 22;273(21):13022-9
references.pmid: 9582338
references.citation: J Biol Chem. 2012 Feb 17;287(8):5492-506
references.pmid: 22194599
references.citation: Matrix Biol. 2014 Apr;35:223-31
references.pmid: 24373744
references.citation: Oncogene. 2001 Nov 1;20(50):7408-12
references.pmid: 11704870
references.citation: J Biol Chem. 2008 Nov 21;283(47):32628-36
references.pmid: 18812315
references.citation: Development. 2003 Apr;130(8):1515-22
references.pmid: 12620978
references.citation: Cancer Res. 2001 Dec 1;61(23):8375-80
references.pmid: 11731411
references.citation: Blood. 2007 Jun 1;109(11):4793-802
references.pmid: 17272502
references.citation: Biochem J. 2011 Nov 15;440(1):157-66
references.pmid: 21871017
references.citation: Matrix Biol. 2014 Apr;35:182-93
references.pmid: 24063949
references.citation: J Biol Chem. 2011 Jan 28;286(4):2636-47
references.pmid: 21075844
references.citation: Mod Pathol. 2010 May;23 Suppl 2:S52-9
references.pmid: 20436503
references.citation: Neuron. 2013 Aug 21;79(4):696-711
references.pmid: 23911103
references.citation: FEBS J. 2013 May;280(10):2271-84
references.pmid: 23374253
references.citation: Oncogene. 2000 Jan 20;19(3):410-6
references.pmid: 10656689
references.citation: Br J Cancer. 2011 Feb 15;104(4):635-42
references.pmid: 21285983
references.citation: Carcinogenesis. 2009 Mar;30(3):397-407
references.pmid: 19126645
references.citation: Nat Commun. 2013;4:1964
references.pmid: 23743995
references.citation: Matrix Biol. 2013 Oct-Nov;32(7-8):393-8
references.pmid: 23727079
references.citation: J Biol Chem. 2006 Sep 8;281(36):26408-18
references.pmid: 16835231
references.citation: Connect Tissue Res. 2008;49(3):230-4
references.pmid: 18661349
references.citation: Int J Biochem Cell Biol. 2003 Mar;35(3):376-90
references.pmid: 12531251
references.citation: Biochim Biophys Acta. 2002 Nov 20;1588(2):165-72
references.pmid: 12385781
references.citation: Dev Biol. 2012 Nov 1;371(1):66-76
references.pmid: 22922164
references.citation: Sci Signal. 2012 Jun 12;5(228):ra42
references.pmid: 22692423
references.citation: Oncogene. 2009 Jan 15;28(2):257-69
references.pmid: 18931701
references.citation: FEBS J. 2010 Oct;277(19):3890-903
references.pmid: 20840586
references.citation: Sci Transl Med. 2013 Apr 10;5(180):180ra48
references.pmid: 23576814
references.citation: Carcinogenesis. 2012 Feb;33(2):326-30
references.pmid: 22159220
references.citation: J Exp Clin Cancer Res. 2004 Dec;23(4):641-50
references.pmid: 15743035
references.citation: Trends Cell Biol. 2010 Jun;20(6):355-62
references.pmid: 20356743
references.citation: FEBS J. 2013 May;280(10):2138-49
references.pmid: 23351020
references.citation: Cancer Res. 2005 Jul 15;65(14):6245-54
references.pmid: 16024626
references.citation: Neuron. 2003 Mar 6;37(5):735-49
references.pmid: 12628165
references.citation: J Biol Chem. 2005 Dec 30;280(52):42573-9
references.pmid: 16253987
references.citation: Am J Pathol. 2012 Aug;181(2):380-7
references.pmid: 22735579
references.citation: Biochem Pharmacol. 2011 May 15;81(10):1171-82
references.pmid: 21371441
references.citation: Int J Cancer. 2008 Dec 1;123(11):2473-9
references.pmid: 18798267
references.citation: EMBO J. 2012 Mar 7;31(5):1095-108
references.pmid: 22343943
references.citation: Tumour Biol. 2015 Mar;36(3):1773-80
references.pmid: 25371074
references.citation: J Biol Chem. 2013 May 17;288(20):13988-99
references.pmid: 23525115
references.citation: Int J Cancer. 2012 May 15;130(10):2271-81
references.pmid: 21671478
references.citation: J Biol Chem. 2003 Feb 28;278(9):7617-23
references.pmid: 12493766
references.citation: J Biol Chem. 2000 Feb 25;275(8):5249-52
references.pmid: 10681494
references.citation: J Cell Biol. 2009 May 18;185(4):743-54
references.pmid: 19433454
references.citation: Matrix Biol. 2013 Jan;32(1):3-13
references.pmid: 23178232
references.citation: J Biol Chem. 1996 May 24;271(21):12532-41
references.pmid: 8647862
references.citation: J Biol Chem. 2010 Dec 31;285(53):42075-85
references.pmid: 20974860
references.citation: Mol Cells. 2004 Apr 30;17(2):181-7
references.pmid: 15179029
references.citation: J Biol Chem. 2014 Jun 6;289(23):16114-28
references.pmid: 24737315
references.citation: Nat Med. 2012 Jun;18(6):883-91
references.pmid: 22635005
references.citation: Dev Cell. 2008 May;14(5):712-25
references.pmid: 18477454
references.citation: EJNMMI Res. 2011 Sep 01;1(1):20
references.pmid: 22214534
references.citation: Cancer Res. 2010 Jun 1;70(11):4453-9
references.pmid: 20460509
references.citation: Connect Tissue Res. 2010 Jun;51(3):230-9
references.pmid: 20073988
references.citation: Matrix Biol. 2013 Oct-Nov;32(7-8):414-23
references.pmid: 23608680
references.citation: Cold Spring Harb Perspect Biol. 2009 Sep;1(3):a002493
references.pmid: 20066107
references.citation: Matrix Biol. 2012 Jan;31(1):3-16
references.pmid: 22033227
references.citation: J Am Soc Mass Spectrom. 2005 Aug;16(8):1221-30
references.pmid: 15979327
references.citation: FEBS J. 2014 Nov;281(22):5023-42
references.pmid: 25333340
references.citation: Front Oncol. 2014 Jan 13;3:327
references.pmid: 24455486
references.citation: Carcinogenesis. 2008 Jul;29(7):1435-40
references.pmid: 18550571
references.citation: J Biol Chem. 2009 Apr 24;284(17):11728-37
references.pmid: 19211552
references.citation: Nature. 2010 Oct 21;467(7318):972-6
references.pmid: 20927103
references.citation: J Exp Med. 2009 Mar 16;206(3):691-705
references.pmid: 19255147
references.citation: Cell Tissue Res. 2010 Jan;339(1):31-46
references.pmid: 19597846
references.citation: Cancer Res. 2001 Jul 15;61(14):5562-9
references.pmid: 11454708
references.citation: Proc Natl Acad Sci U S A. 2013 Apr 16;110(16):6400-5
references.pmid: 23509287
references.citation: J Biol Chem. 2006 Nov 17;281(46):35116-28
references.pmid: 16870619
references.citation: Breast Cancer Res Treat. 2006 Jul;98(1):91-8
references.pmid: 16636895
references.citation: Autophagy. 2010 Nov;6(8):1090-106
references.pmid: 20890124
references.citation: Nature. 2009 Jan 1;457(7225):102-6
references.pmid: 19122641
references.citation: J Cell Sci. 2010 Nov 1;123(Pt 21):3796-807
references.pmid: 20971705
references.citation: Matrix Biol. 2010 Jul;29(6):442-52
references.pmid: 20416374
references.citation: Mol Cell Biol. 2010 Apr;30(7):1634-49
references.pmid: 20100867
references.citation: Tumour Biol. 2014 Oct;35(10):9649-59
references.pmid: 24969560
references.citation: J Cell Sci. 2010 May 15;123(Pt 10):1603-11
references.pmid: 20445011
references.citation: IUBMB Life. 2006 Jan;58(1):47-53
references.pmid: 16540432
references.citation: Cancer Metastasis Rev. 2011 Jun;30(2):253-68
references.pmid: 21308479
references.citation: Cancer Res. 2005 Jul 1;65(13):5778-84
references.pmid: 15994953
references.citation: Cancer Res. 2011 Feb 1;71(3):645-54
references.pmid: 21266359
references.citation: Dis Model Mech. 2013 Jan;6(1):25-39
references.pmid: 23268536
references.citation: EMBO J. 2002 Dec 2;21(23):6473-82
references.pmid: 12456654
references.citation: J Histochem Cytochem. 2012 Dec;60(12):963-75
references.pmid: 22821552
references.citation: Nat Cell Biol. 2014 Dec;16(12):1238-48
references.pmid: 25362351
references.citation: Science. 2012 Nov 16;338(6109):889-90
references.pmid: 23161981
references.citation: Br J Cancer. 2008 Jun 17;98(12):1993-8
references.pmid: 18542065
references.citation: EMBO Rep. 2010 Nov;11(11):854-60
references.pmid: 20930847
references.citation: J Immunol. 2004 May 1;172(9):5185-93
references.pmid: 15100255
references.citation: FEBS J. 2013 May;280(10):2294-306
references.pmid: 23374281
references.citation: J Biol Chem. 2003 Jul 11;278(28):26227-37
references.pmid: 12719420
references.citation: J Biol Chem. 2013 Apr 5;288(14):10093-9
references.pmid: 23430739
references.citation: Clin Cancer Res. 2007 Feb 1;13(3):806-15
references.pmid: 17289871
references.citation: Nat Cell Biol. 2012 Jul;14(7):677-85
references.pmid: 22660413
references.citation: Proc Natl Acad Sci U S A. 2011 Oct 11;108(41):17022-7
references.pmid: 21969569
references.citation: Int J Biochem Cell Biol. 2014 Sep;54:223-35
references.pmid: 25091702
references.citation: BMC Cancer. 2011;11:314
references.pmid: 21791066
references.citation: PLoS One. 2013;8(12):e85737
references.pmid: 24392029
references.citation: Eur J Immunol. 2011 Feb;41(2):437-49
references.pmid: 21268013
references.citation: Nature. 2007 Oct 11;449(7163):682-8
references.pmid: 17898713
references.citation: J Biol Chem. 1993 May 25;268(15):11440-8
references.pmid: 8496192
references.citation: Dev Cell. 2011 Jul 19;21(1):77-91
references.pmid: 21763610
references.citation: Nat Genet. 2003 Dec;35(4):367-71
references.pmid: 14625552
references.citation: Matrix Biol. 2014 Apr;35:132-42
references.pmid: 24361484
references.citation: J Biol Chem. 2004 May 28;279(22):23536-41
references.pmid: 15044433
references.citation: FASEB J. 2003 Mar;17(3):464-6
references.pmid: 12631584
references.citation: Proc Natl Acad Sci U S A. 2013 Jul 9;110(28):E2582-91
references.pmid: 23798385
references.citation: Biochim Biophys Acta. 2014 Aug;1840(8):2460-70
references.pmid: 24508120
references.citation: Circ Res. 2004 Apr 30;94(8):1067-74
references.pmid: 15031262
references.citation: Dev Biol. 2010 Dec 1;348(1):107-18
references.pmid: 20883685
references.citation: FASEB J. 2014 Jun;28(6):2455-65
references.pmid: 24558194
references.citation: Atherosclerosis. 2001 Mar;155(1):87-97
references.pmid: 11223430
references.citation: Development. 2005 Feb;132(4):659-66
references.pmid: 15647318
references.citation: PLoS One. 2012;7(2):e30992
references.pmid: 22363530
references.citation: J Histochem Cytochem. 2011 Jun;59(6):615-29
references.pmid: 21430259
references.citation: Int J Clin Exp Pathol. 2013;6(11):2497-505
references.pmid: 24228112
references.citation: FEBS J. 2013 May;280(10):2353-68
references.pmid: 23350987
references.citation: J Biol Chem. 2014 Feb 28;289(9):5499-509
references.pmid: 24403068
references.citation: Biochem Biophys Res Commun. 1993 Jan 29;190(2):616-23
references.pmid: 8427603
references.citation: Curr Opin Cell Biol. 2009 Oct;21(5):662-9
references.pmid: 19535238
references.citation: Oncotarget. 2014 Apr 15;5(7):1885-96
references.pmid: 24681892
references.citation: Br J Cancer. 2007 May 21;96(10):1560-8
references.pmid: 17453002
references.citation: Oncogene. 1996 Oct 17;13(8):1789-99
references.pmid: 8895526
references.citation: J Biol Chem. 2004 Oct 15;279(42):44084-92
references.pmid: 15292202
references.citation: Cells. 2012 Jul 26;1(3):325-45
references.pmid: 24710479
references.citation: Mol Cell Proteomics. 2012 Jun;11(6):M111.011403
references.pmid: 22318369
references.citation: Anticancer Res. 2006 Jan-Feb;26(1B):621-7
references.pmid: 16739330
references.citation: Nat Cell Biol. 2010 Feb;12(2):119-31
references.pmid: 20098416
references.citation: Curr Biol. 2004 Oct 26;14(20):1852-7
references.pmid: 15498494
references.citation: Cell Oncol. 2009;31(1):19-29
references.pmid: 19096147
references.citation: J Biol Chem. 2011 Jul 22;286(29):25947-62
references.pmid: 21596751
references.citation: J Biochem. 2013 Jul;154(1):41-50
references.pmid: 23576453
references.citation: Cancer Res. 2012 Jan 1;72(1):112-21
references.pmid: 22094876
references.citation: Cancer Res. 2006 Feb 1;66(3):1455-63
references.pmid: 16452201
references.citation: PLoS One. 2010;5(11):e13828
references.pmid: 21079779
references.citation: Annu Rev Biochem. 1998;67:609-52
references.pmid: 9759499
references.citation: J Histochem Cytochem. 2012 Dec;60(12):898-907
references.pmid: 23042481
references.citation: J Cell Biol. 2003 Nov 10;163(3):625-35
references.pmid: 14610063
references.citation: Hum Mol Genet. 2011 Dec 15;20(24):4891-902
references.pmid: 21920942
references.citation: Exp Biol Med (Maywood). 2011 Nov;236(11):1247-53
references.pmid: 21998129
references.citation: Exp Cell Res. 2003 Jun 10;286(2):219-32
references.pmid: 12749851
references.citation: FEBS J. 2010 Oct;277(19):3904-23
references.pmid: 20840587
references.citation: Cancer Res. 2005 May 15;65(10):4442-7
references.pmid: 15899837
references.citation: Cancer Res. 2005 Mar 1;65(5):1748-54
references.pmid: 15753370
references.citation: Front Oncol. 2014 Jul 24;4:195
references.pmid: 25105093
references.citation: Nature. 2012 Jun 21;486(7403):410-4
references.pmid: 22722203
references.citation: Oncogene. 2005 Feb 3;24(6):1104-10
references.pmid: 15690056
references.citation: Blood. 2010 Mar 25;115(12):2449-57
references.pmid: 20097882
references.citation: Matrix Biol. 2013 Jun 24;32(5):223-7
references.pmid: 23499529
references.citation: Oncology. 2004;67(1):11-8
references.pmid: 15459490
references.citation: Clin Exp Metastasis. 2011 Feb;28(2):113-25
references.pmid: 21153687
references.citation: J Biol Chem. 2009 Sep 4;284(36):24035-48
references.pmid: 19605353
references.citation: FEBS J. 2013 May;280(10):2320-31
references.pmid: 23384311
references.citation: J Biol Chem. 1997 Mar 28;272(13):8133-6
references.pmid: 9079625
references.citation: Dev Cell. 2010 Apr 20;18(4):605-20
references.pmid: 20412775
references.citation: PLoS One. 2013;8(11):e80084
references.pmid: 24223213
references.citation: Matrix Biol. 2014 Apr;35:103-11
references.pmid: 24447998
references.citation: J Biol Chem. 2000 Oct 20;275(42):32879-87
references.pmid: 10913155
references.citation: Clin Cancer Res. 2009 Jan 15;15(2):731-9
references.pmid: 19147781
references.citation: Dev Cell. 2008 May;14(5):700-11
references.pmid: 18477453
references.citation: Nat Cell Biol. 2011 Feb;13(2):132-41
references.pmid: 21258367
references.citation: Am J Pathol. 2014 Jan;184(1):28-41
references.pmid: 24269592
references.citation: Biochim Biophys Acta. 2012 Dec;1820(12):1926-39
references.pmid: 22884656
references.citation: Breast Cancer Res Treat. 2014 Sep;147(2):423-31
references.pmid: 25007959
references.citation: J Cell Biol. 2013 Feb 18;200(4):367-71
references.pmid: 23420870
references.citation: PLoS One. 2012;7(9):e45559
references.pmid: 23029096
references.citation: Arch Oral Biol. 2012 May;57(5):531-8
references.pmid: 22134060
references.citation: Tumour Biol. 2012 Jun;33(3):679-88
references.pmid: 22389160
references.citation: Breast Cancer Res Treat. 2003 Jul;80(2):221-32
references.pmid: 12908826
references.citation: Dev Cell. 2007 Jul;13(1):57-71
references.pmid: 17609110
references.citation: Biochemistry. 1999 Sep 14;38(37):12187-95
references.pmid: 10508424
references.citation: Leuk Lymphoma. 2012 Sep;53(9):1764-8
references.pmid: 22335531
references.citation: Matrix Biol. 2014 Feb;34:46-54
references.pmid: 24472739
references.citation: Development. 2010 Jun;137(12):2033-44
references.pmid: 20501592
references.citation: Genome Biol. 2008;9(5):224
references.pmid: 18505598
references.citation: Curr Opin Cell Biol. 2010 Apr;22(2):124-31
references.pmid: 20034776
references.citation: Oncogene. 2003 Dec 18;22(58):9243-53
references.pmid: 14681683
references.citation: Curr Opin Genet Dev. 2012 Feb;22(1):56-7
references.pmid: 22326829
references.citation: Biochim Biophys Acta. 2006 Aug;1760(8):1217-25
references.pmid: 16730906
references.citation: Science. 2003 Mar 28;299(5615):2039-45
references.pmid: 12663920
references.citation: Proc Natl Acad Sci U S A. 2013 Oct 22;110(43):17380-5
references.pmid: 24101524
references.citation: Nature. 1999 Jul 15;400(6741):276-80
references.pmid: 10421371
references.citation: Matrix Biol. 2014 Apr;35:266-75
references.pmid: 24373743
references.citation: Breast Cancer. 2012 Jan;19(1):46-53
references.pmid: 21505857
references.citation: Tumour Biol. 2015 Mar;36(3):1731-8
references.pmid: 25361632
references.citation: J Cell Biol. 2011 Feb 21;192(4):691-704
references.pmid: 21339334
references.citation: Genes Cells. 2001 Mar;6(3):237-47
references.pmid: 11260267
references.citation: BMC Cancer. 2013;13:517
references.pmid: 24180670
references.citation: J Biol Chem. 2005 Jan 21;280(3):2116-25
references.pmid: 15537637
references.citation: J Biol Chem. 2012 Dec 21;287(52):43543-56
references.pmid: 23060442
references.citation: Cell Tissue Res. 1992 Apr;268(1):167-77
references.pmid: 1499048
references.citation: Dev Cell. 2009 Oct;17(4):470-81
references.pmid: 19853561
references.citation: Cancer Biol Ther. 2007 Apr;6(4):587-95
references.pmid: 17457040
references.citation: Histopathology. 2006 Dec;49(6):582-93
references.pmid: 17163843
references.citation: Mol Biol Cell. 2010 Nov 15;21(22):4028-41
references.pmid: 20861306
references.citation: J Biomech Eng. 2005 Feb;127(1):181-5
references.pmid: 15868800
references.citation: Blood. 2012 Feb 16;119(7):1683-92
references.pmid: 22117039
references.citation: J Cell Mol Med. 2009 Sep;13(9B):3687-98
references.pmid: 19298526
references.citation: Cancer. 2003 Aug 1;98(3):474-83
references.pmid: 12879463
references.citation: Gastroenterology. 2001 Sep;121(3):657-67
references.pmid: 11522750
references.citation: PLoS One. 2014;9(4):e95240
references.pmid: 24740415
references.citation: Cell Prolif. 2011 Aug;44(4):343-51
references.pmid: 21702857
references.citation: J Biol Chem. 2012 Sep 21;287(39):32298-306
references.pmid: 22859297
references.citation: Matrix Biol. 2013 Aug 8;32(6):289-97
references.pmid: 23583522
references.citation: Oncogene. 2002 May 23;21(23):3688-95
references.pmid: 12032837
references.citation: Annu Rev Cell Dev Biol. 2010;26:89-114
references.pmid: 20565253
references.citation: Trends Mol Med. 2013 Jul;19(7):410-7
references.pmid: 23639582
references.citation: Anticancer Res. 2007 Sep-Oct;27(5A):3045-50
references.pmid: 17970043
references.citation: Eur J Biochem. 2003 Oct;270(19):3996-4007
references.pmid: 14511382
references.citation: Eur J Biochem. 1995 Oct 1;233(1):192-9
references.pmid: 7588746
references.citation: Br J Cancer. 2012 Mar 13;106(6):1214-23
references.pmid: 22374465
references.citation: Surgery. 2002 Aug;132(2):326-33
references.pmid: 12219030
references.citation: Gene Ther. 2015 Mar;22(3):247-56
references.pmid: 25503693
references.citation: Dev Cell. 2001 Aug;1(2):251-64
references.pmid: 11702784
references.citation: PLoS One. 2011;6(6):e21106
references.pmid: 21698156
references.citation: Int J Cancer. 2012 Sep 15;131(6):E884-96
references.pmid: 22573479
references.citation: BMC Cancer. 2013;13:24
references.pmid: 23327652
references.citation: Breast Cancer Res. 2011;13(3):R58
references.pmid: 21658254
references.citation: Arch Gynecol Obstet. 2014 Mar;289(3):649-55
references.pmid: 24013431
references.citation: Cancer Cell. 2013 Mar 18;23(3):287-301
references.pmid: 23416000
references.citation: Cancer Cell. 2008 Jun;13(6):472-82
references.pmid: 18538731
references.citation: Cancer Cell. 2012 Mar 20;21(3):297-308
references.pmid: 22439925
references.citation: Arch Gynecol Obstet. 2014 Jun;289(6):1315-23
references.pmid: 24385285
references.citation: BMC Cancer. 2013;13:34
references.pmid: 23351331
references.citation: Clin Cancer Res. 2002 Apr;8(4):1054-60
references.pmid: 11948113
references.citation: FEBS J. 2010 Oct;277(19):3876-89
references.pmid: 20840585
references.citation: N Engl J Med. 2013 Feb 14;368(7):651-62
references.pmid: 23406030
references.citation: Mol Oncol. 2008 Oct;2(3):223-32
references.pmid: 19383343
references.citation: Dev Neurobiol. 2011 Jul;71(7):608-18
references.pmid: 21500363
references.citation: J Clin Invest. 2005 Aug;115(8):2223-33
references.pmid: 16025156
references.citation: Dev Cell. 2004 Oct;7(4):503-12
references.pmid: 15469839
references.citation: Clin Exp Med. 2012 Sep;12(3):195-9
references.pmid: 21879307
references.citation: PLoS One. 2012;7(12):e52175
references.pmid: 23300607
references.citation: J Biol Chem. 2007 May 4;282(18):13326-33
references.pmid: 17347152
references.citation: Thromb Haemost. 2008 Jan;99(1):133-41
references.pmid: 18217145
references.citation: Hepatology. 2014 Mar;59(3):958-73
references.pmid: 24005989
references.citation: Anticancer Res. 2002 Sep-Oct;22(5):2893-8
references.pmid: 12530013
references.citation: Expert Opin Ther Targets. 2013 Jun;17(6):695-705
references.pmid: 23421390
references.citation: Matrix Biol. 2014 Apr;35:8-13
references.pmid: 24134926
references.citation: Br J Cancer. 2010 Apr 13;102(8):1265-75
references.pmid: 20332776
references.citation: Biochem J. 2008 Mar 15;410(3):503-11
references.pmid: 17967162
references.citation: Science. 2011 Dec 16;334(6062):1573-7
references.pmid: 22174255
references.citation: J Am Coll Surg. 2005 Mar;200(3):328-35
references.pmid: 15737842
references.citation: Development. 2003 Dec;130(25):6245-55
references.pmid: 14602684
references.citation: Science. 2012 Nov 16;338(6109):956-9
references.pmid: 23112296
references.citation: J Biol Chem. 2002 Jan 4;277(1):623-9
references.pmid: 11679586
references.citation: Nat Cell Biol. 2010 Jan;12(1):19-30; sup pp 1-13
references.pmid: 19966785
references.citation: J Cell Biol. 2004 Oct 11;167(1):171-81
references.pmid: 15479743
references.citation: Cancer Res. 2006 Nov 1;66(21):10233-7
references.pmid: 17079438
references.citation: PLoS One. 2013;8(2):e56616
references.pmid: 23424670
references.citation: Proc Natl Acad Sci U S A. 1999 Mar 16;96(6):3092-7
references.pmid: 10077642
references.citation: Science. 2011 Jun 17;332(6036):1429-33
references.pmid: 21617040
references.citation: Am J Clin Pathol. 1999 Sep;112(3):377-83
references.pmid: 10478144
references.citation: Proc Natl Acad Sci U S A. 1992 Feb 1;89(3):932-6
references.pmid: 1736308
references.citation: Annu Rev Biochem. 2014;83:129-57
references.pmid: 24606135
references.citation: J Biol Chem. 2007 May 18;282(20):14906-15
references.pmid: 17344212
references.citation: Sci Signal. 2011;4(199):ra75
references.pmid: 22087031
references.citation: J Oral Pathol Med. 2010 Mar;39(3):230-5
references.pmid: 20070486
references.citation: PLoS One. 2011;6(5):e19771
references.pmid: 21573075
references.citation: Mol Cancer Res. 2013 May;11(5):443-55
references.pmid: 23449934
references.citation: J Hepatol. 2008 Aug;49(2):207-16
references.pmid: 18490072
references.citation: Clin Biochem. 2013 Jul;46(10-11):926-32
references.pmid: 23415719
references.citation: J Biol Chem. 2009 May 15;284(20):13843-55
references.pmid: 19279012
references.citation: Curr Biol. 2005 May 10;15(9):833-8
references.pmid: 15886101
references.citation: PLoS One. 2012;7(4):e35128
references.pmid: 22514714
references.citation: Nat Genet. 1998 Sep;20(1):78-82
references.pmid: 9731537
references.citation: PLoS One. 2013;8(10):e78157
references.pmid: 24205138
references.citation: Int J Cancer. 2010 Feb 1;126(3):640-50
references.pmid: 19662655
references.citation: Exp Cell Res. 2004 Oct 15;300(1):234-47
references.pmid: 15383330
references.citation: Cold Spring Harb Perspect Biol. 2012 Nov;4(11). pii: a011338. doi: 10.1101/cshperspect.a011338
references.pmid: 23125018
references.citation: Clin Dev Immunol. 2011;2011:842849
references.pmid: 22190973
references.citation: FEBS J. 2011 Oct;278(19):3782-92
references.pmid: 21834878
references.citation: Cell. 2008 Jun 27;133(7):1214-27
references.pmid: 18585355
references.citation: PLoS One. 2010;5(11):e15394
references.pmid: 21072212
references.citation: Blood. 2012 Feb 2;119(5):1292-301
references.pmid: 22144179
references.citation: J Mol Biol. 2000 Dec 8;304(4):669-80
references.pmid: 11099388
references.citation: J Clin Invest. 2010 Dec;120(12):4251-72
references.pmid: 21084753
references.citation: Nat Rev Mol Cell Biol. 2012 Dec;13(12):780-8
references.pmid: 23175281
references.citation: Autophagy. 2011 Mar;7(3):297-300
references.pmid: 21252623
references.citation: J Biol Chem. 2014 Dec 19;289(51):35237-45
references.pmid: 25391648
references.citation: Oncogene. 2006 Mar 2;25(9):1408-12
references.pmid: 16247452
references.citation: J Natl Cancer Inst. 2004 Aug 18;96(16):1219-30
references.pmid: 15316057
references.citation: Nat Rev Mol Cell Biol. 2007 Dec;8(12):957-69
references.pmid: 17971838
references.citation: FEBS J. 2013 May;280(10):2207-15
references.pmid: 23375101
references.citation: Matrix Biol. 2014 Apr;35:82-90
references.pmid: 24389353
references.citation: FEBS J. 2012 Apr;279(7):1177-97
references.pmid: 22333131
references.citation: Cancer Res. 2013 Aug 15;73(16):5016-28
references.pmid: 23824740
references.citation: Stem Cells. 2012 Sep;30(9):1863-74
references.pmid: 22761013
references.citation: FEBS J. 2013 May;280(10):2248-59
references.pmid: 23374155
references.citation: J Biol Chem. 2010 Mar 12;285(11):8472-80
references.pmid: 20064927
references.citation: Funct Integr Genomics. 2011 Jun;11(2):215-24
references.pmid: 21061036
references.citation: J Biol Chem. 2014 Feb 21;289(8):4952-68
references.pmid: 24403067
references.citation: Cell. 2008 Jan 11;132(1):27-42
references.pmid: 18191218
references.citation: Am J Pathol. 2008 Sep;173(3):844-55
references.pmid: 18688028
references.citation: Mol Cancer Res. 2012 Jun;10(6):689-702
references.pmid: 22513363
references.citation: Am J Pathol. 2011 Mar;178(3):1221-32
references.pmid: 21356373
references.citation: J Cell Mol Med. 2011 May;15(5):1013-31
references.pmid: 21155971
references.citation: Curr Chem Biol. 2010 Jan 1;4(1):20-31
references.pmid: 20596243
references.citation: Nat Genet. 1996 Feb;12(2):186-90
references.pmid: 8563758
references.citation: J Biol Chem. 2008 Oct 24;283(43):28881-7
references.pmid: 18708345
references.citation: J Clin Invest. 2011 Oct;121(10):3786-8
references.pmid: 21965334
references.citation: J Biol Chem. 2012 Jul 13;287(29):24320-9
references.pmid: 22582394
references.citation: FEBS J. 2013 May;280(10):2477-89
references.pmid: 23374223
references.citation: J Bone Miner Res. 2010 Jun;25(6):1295-304
references.pmid: 20200931
references.citation: Nat Genet. 2011 May;43(5):459-63
references.pmid: 21441931
references.citation: Am J Pathol. 2011 Jan;178(1):325-35
references.pmid: 21224069
references.citation: BMC Cancer. 2012;12:341
references.pmid: 22862967
references.citation: Biomed Res Int. 2013;2013:852093
references.pmid: 23984412
references.citation: Curr Med Chem. 2011;18(11):1584-98
references.pmid: 21428887
references.citation: Cardiovasc Res. 2008 Jul 15;79(2):208-17
references.pmid: 18430751
references.citation: Clin Cancer Res. 2003 Jan;9(1):207-14
references.pmid: 12538471
references.citation: Integr Biol (Camb). 2013 Aug;5(8):1045-56
references.pmid: 23817524
references.citation: Biochim Biophys Acta. 2014 Aug;1840(8):2482-90
references.pmid: 24447566
references.citation: J Biol Chem. 2011 Feb 25;286(8):6490-9
references.pmid: 21131364
references.citation: Chembiochem. 2008 Jan 25;9(2):198-200
references.pmid: 18085541
references.citation: J Biol Chem. 2012 Apr 20;287(17):14040-51
references.pmid: 22351761
references.citation: Matrix Biol. 2014 Apr;35:194-205
references.pmid: 24361483
references.citation: Genet Mol Res. 2011;10(4):3901-13
references.pmid: 22183949
references.citation: Med Oncol. 2013 Dec;30(4):707
references.pmid: 23996242
references.citation: J Biol Chem. 2001 Nov 16;276(46):42774-81
references.pmid: 11533057
references.citation: Antioxid Redox Signal. 2011 Jun;14(11):2251-69
references.pmid: 20712403
references.citation: Am J Pathol. 2014 Aug;184(8):2146-53
references.pmid: 24976620
references.citation: Nat Rev Mol Cell Biol. 2007 Sep;8(9):741-52
references.pmid: 17717517
references.citation: Matrix Biol. 2014 Apr;35:91-102
references.pmid: 24447999
references.citation: Cancer Res. 2011 Apr 15;71(8):3162-72
references.pmid: 21289131
references.citation: Am J Pathol. 2002 Jan;160(1):185-94
references.pmid: 11786412
references.citation: J Theor Biol. 1982 Nov 7;99(1):31-68
references.pmid: 6892044
references.citation: Cancer Cell. 2004 Jul;6(1):17-32
references.pmid: 15261139
references.citation: FEBS J. 2013 Jun;280(12):2870-87
references.pmid: 23601700
references.citation: FEBS J. 2013 May;280(10):2342-52
references.pmid: 23387827
references.citation: Mol Cancer. 2015;14:15
references.pmid: 25623282
references.citation: Cell. 2004 Oct 15;119(2):231-44
references.pmid: 15479640
references.citation: Matrix Biol. 2014 Apr;35:143-51
references.pmid: 24480070
references.citation: Clin Cancer Res. 2004 Apr 1;10(7):2491-8
references.pmid: 15073129
references.citation: OMICS. 2009 Aug;13(4):345-54
references.pmid: 19422301
references.citation: Tumour Biol. 2013 Feb;34(1):131-7
references.pmid: 23007878
references.citation: PLoS One. 2011;6(11):e26396
references.pmid: 22096483
references.citation: Autophagy. 2013 Oct;9(10):1626-8
references.pmid: 23989617
references.citation: Clin Cancer Res. 2011 Mar 15;17(6):1382-93
references.pmid: 21257720